Methylome based analysis and treatment for meningioma

ABSTRACT

There is described herein a method of predicting recurrence free survival in a patient with meningioma comprising: (a) determining a tumor DNA methylation profile from a tumor sample from the patient, the tumor DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9; and (b) calculating a risk of meningioma recurrence based on comparing the tumor DNA methylation profile and a reference methylation profile comprising the extent to which the methylation status of the at least 200 loci is associated with a risk of recurrence.

RELATED APPLICATIONS

This application claims priority to U.S. Provisional Application No. 62/693,928, which is hereby incorporated by reference in its entirety.

FIELD OF THE INVENTION

The invention relates to compositions and methods for use in treating and predicting meningioma recurrence.

BACKGROUND OF THE INVENTION

Meningiomas are the most common primary intracranial tumor. They account for 37% of all central nervous system tumors, and are continuing to increase in incidence with the ageing population¹. They result in significant neurological morbidity and loss of quality of life by exerting mass effect on critical adjacent brain regions¹. The current standard-of-care for nearly all patients with symptomatic meningiomas includes gross total tumor resection with removal of involved dura and bone when possible². However, despite radical surgical resection, approximately 20% of meningiomas display aggressive behaviour with early tumor recurrence resulting in a clinical course of repetitive disease- and treatment-related morbidity. Radiation therapy can be used to provide disease control as an adjunct to surgery for a subset of tumors^(3,4). However, radiotherapy can often result in adverse radiation effects that lead to considerable morbidity and neurological dysfunction long-term precluding universal use in all patients⁵. One of the greatest clinical challenges faced by clinicians is the inability to predict early tumor recurrence at an individual patient level which limits the appropriate selection of patients who would benefit from adjuvant radiation therapy.

To date, the most reliable clinical factors associated with recurrence in meningiomas have been the World Health Organization (WHO) grade of the tumor and extent of tumor resection at surgery^(2,6). Although both are crudely associated with recurrence rates on a population level, they are challenged with inter-rater variability of grading and considerable within-grade variation of recurrence risk for individual patients^(2,6). In the past decade, several studies have focused on molecular profiling of meningiomas to refine biological subgroups⁷⁻¹³. With the exception of mutations in BAP1 and TERT promoter, each of which occur rarely in these tumors, the mutations identified in meningioma have not been shown to be tightly correlated to patient outcome with current standard of care⁷⁻¹⁰. We and others have independently shown that global DNA methylation profiling reveals robust methylome-based meningioma subtypes, however, the clinical translation of this to predict recurrence risk for individual patients has not been demonstrated to date.

SUMMARY OF THE INVENTION

To examine whether methylation profiles can be defined and validated for clinical utility, we aimed to develop and validate a methylome-based predictor of early meningioma recurrence that could be combined with established prognostic clinical factors to individualize decisions regarding the need for post-operative therapeutic interventions, in particular, whether to treat patients with adjuvant radiation therapy versus observation alone. Our work is the first in neuro-oncology to demonstrate the transformative utility of integrating clinical and molecular factors for use beyond simple classification into the realm of individualized prognostication.

Our work is the first to demonstrate the transformative utility of integrating clinical and molecular factors for use beyond simple classification into the realm of individualized prognostication for any brain tumor. Using our developed and validated tools that are publicly available, clinicians will be able combine clinical and molecular factors to determine an individualized probability of recurrence for patients with meningiomas. This represents a major advance in the field of personalized medicine for neuro-oncology, and the use of this tool can help clinicians overcome one of the most challenging limitations we face when treating patients with meningiomas.

In an aspect, there is provided a method of predicting recurrence free survival in a patient with meningioma comprising:

-   -   (a) determining a tumor DNA methylation profile from a tumor         sample from the patient, the tumor DNA methylation profile         comprising the methylation status of at least 200 loci         represented by the probes set forth in Table 9;     -   (b) calculating a risk of meningioma recurrence based on         comparing the tumor DNA methylation profile and a reference         methylation profile comprising the extent to which the         methylation status of the at least 200 loci is associated with a         risk of recurrence.

In an aspect, there is provided an array consisting of the 9529 probes set forth in Table 9.

In an aspect, there is provided a computer program product for use in conjunction with a computer having a processor and a memory connected to the processor, the computer program product comprising a computer readable storage medium having a computer mechanism encoded thereon, wherein the computer program mechanism may be loaded into the memory of the computer and cause the computer to carry out the method described herein.

In an aspect, there is provided a computer implemented product for predicting recurrence free survival in a patient with meningioma comprising:

-   -   (a) a means for receiving values corresponding to a tumor DNA         methylation profile from a tumor sample from the patient, the         DNA methylation profile comprising the methylation status of at         least 200 loci represented by the probes set forth in Table 9;     -   (b) a database comprising a reference methylation profile         representing the extent to which each said methylation status of         the at least 200 loci is associated with a risk of recurrence;         and     -   (c) a means for calculating a risk of meningioma recurrence         based on a comparison of the tumor DNA methylation profile and         the reference methylation profile.

In an aspect, there is provided a computer readable medium having stored thereon a data structure for storing the computer implemented product described herein. In an aspect, there is provided a computer system comprising

-   -   (a) a user interface capable of receiving values corresponding         to a tumor DNA methylation profile from a tumor sample from the         patient, the DNA methylation profile comprising the methylation         status of at least 200 loci represented by the probes set forth         in Table 9;     -   (b) a database including records comprising a reference         methylation profile comprising the extent to which each said         methylation status of the at least 200 loci is associated with a         risk of recurrence;     -   (c) an output that displays a prediction of recurrence free         survival based on a comparison of the tumor DNA methylation         profile and the reference methylation profile. In an aspect,         there is provided a method of predicting recurrence free         survival in a patient with meningioma comprising:     -   (a) determining a tumor DNA methylation profile from a tumor         sample from the patient, the tumor DNA methylation profile         comprising the methylation status of at least 200 genes set         forth in Table 9;     -   (b) calculating a risk of meningioma recurrence based on         comparing the tumor DNA methylation profile and a reference         methylation profile setting forth the extent to which the         methylation status of the at least 200 genes is associated with         a risk of recurrence.

BRIEF DESCRIPTION OF FIGURES

These and other features of the preferred embodiments of the invention will become more apparent in the following detailed description in which reference is made to the appended drawings wherein:

FIG. 1 shows comparison of grade-based and methylome-based recurrence-free survival predictor performance. Data presented are time-dependent receiver operating characteristic curve (ROC) and average area under the curve (AUC) as well as ΔAUC (DAUC) with 95% CI using 10,000 bootstrap resampling validation approach for methylome-based and grade-based predictors in the (A) first validation cohort and (B) second validation cohort (C) third validation cohort, (D) combined validation cohorts.

FIG. 2 shows recurrence-free survival (RFS) analysis of the first validation cohort (A), second validation cohort (B), and (C) third validation cohort using the 5y methylome-based RFS predictor, based on separation into distinct risk groups by median.

FIG. 3 shows frequency of copy number alterations across the genome stratified in risk groups according to the methylome-based predictor. Groups correspond to same groups seen in FIG. 2.

FIG. 4 shows recurrence-free survival (RFS) analysis of all WHO grade I (A), WHO grade II (B) and WHO grade III (C) tumors in all three validation cohorts.

FIG. 5 shows a nomogram to predict 5-year recurrence risk in meningiomas (A) Total points generated from scoring of methylome-based RFS predictor, WHO grade, and Simpson grade are tallied in the calculator and correlated to 5-year RFS in the nomogram in the Risk Nomogram. (B) Time-dependent average AUC with 95% CI as well as ΔAUC (DAUC) with 95% CI using 10,000 bootstrap resampling validation approach generated for the meningioma recurrence score and a nomogram using clinical factors alone in the first and third validation cohorts as well as both combined validation cohorts. (C) Calibration curve of the nomogram to predict recurrence-free survival at 5 years in the combined validation cohort. The observed recurrence-free survival is plotted on the y-axis and nomogram predicted probability is plotted on the x-axis.

FIG. 6 shows the meningioma recurrence score identifies two risk groups (high risk versus low risk) that may help individualize adjuvant management decisions such as the need for radiation therapy in patients with meningiomas. Patient tumor samples can be interrogated for DNA methylation profile of selected 9529 probes and a 5-year methylome-based RFS predictor score is generated. This score is combined with tumor WHO grade and Simpson grade in the meningioma recurrence score to develop an individualized probability of 5-year recurrence-free survival.

FIG. 7 shows outline of the overall study design. Illumina Infinium DNA methylation arrays were generated on meningioma samples (from Toronto), and additional methylation data from published work (Europe, Houston) were used to develop and validate the methylome-based predictor for recurrence-free survival (RFS). Cox modeling of individual probes was use for feature selection in a training set (N=228 patients) which was then applied to 3 independent validation sets (N=54, N=140, and N=64 patients). Genome-wide copy number alterations were inferred from DNA methylation data. Functional annotation of genes recurrently represented by the selected probes was performed and gene-expression analysis was correlated to DNA methylation profiles using two publicly available microarray datasets. Cox modeling was used to generate a 5-year meningioma recurrence score based on a nomogram that integrated the validated methylome-based predictor with established key clinical factors.

FIG. 8 shows sources of variability in feature selection from (A,C) 71538 probes that were highly correlated with RFS (P<0.01) in training set (B,D) 23571 probes common in samples from training set (University Health Network and University of Heidelberg FFPE and fresh frozen). Results demonstrate that with 23571 probes common in samples in the training set tissue source was not a major source of variation with convergence between the center of gravity and standard deviation on principal component analysis.

FIG. 9 shows enrichment scores from functional clustering of genes with minimum 5 annotated probes (* indicates Bonferonni adjusted p-values<0.05).

FIG. 10 shows nomogram calculator and risk nomogram. For a given patient, the cumulative points from each variable (“5y methylome probability”, “WHO grade” and “Simpson grade”) is calculated for a result of total points. The “risk nomogram” is then used to scale the total points to risk of 5-year RFS. A patient with a WHO grade 2 meningioma (23 points), simpson grade 3 resection (10 points), and 5-year methylome probability score of 0.6 (60 points) would have a cumulative 94 points, which would result in a 5-year predicted RFS probability of 40.5%. The 5-year methylome probability score would be generated via an online platform that utilizes the generated and validated predictor to create probability scores from methylation array-based analysis of a tumor sample (IDAT file).

FIG. 11 shows plot of martingale residuals versus methylation-based predictor demonstrating appropriateness to model methylation-based predictor as a linear function in Cox model.

FIG. 12 shows plot of Schoenfeld residuals versus time for each covariate included in main Cox model demonstrating no violation of proportional hazard assumption.

FIG. 13 shows internal validation using 10,000 bootstramp resampling method and and time-dependent AUC calculations of meningioma recurrence score. Solid lines represents the mean AUC, darkened grey area represents the interquartile range, and the light grey area represents the minimum and maximum.

FIG. 14 shows case example #1 demonstrating power of individualized prognostication. This is a case of a 58-year-old male with a right sided sphenoid wing meningioma that was resected completely (simpson grade 1). Histopathology demonstrated a WHO Grade I meningioma, 1 mitosis per 10HPF, no necrosis or other atypical features. Traditionally this patient would be considered cured of their disease (benign meningioma that is completely resected), however, in fact this patient experienced 30-months after initial resection. The methylation-based predictor score for this patient is 0.9785 (score ranges from 0 [lowest risk] to 1 [highest risk]). (A) The meningioma recurrence score for this patient demonstrates a 71.75% risk of recurrence within 5-years making this patient high risk for recurrence. (B) Copy number variation plot demonstrates a bland copy number landscape and does not demonstrate a high degree of burden of copy number alterations. (C) Existing DNA methylation-classification systems deem this tumor to be of intermediate risk without clear predictions of 5-year recurrence risk.

FIG. 15 shows Case example #2 demonstrating power of individualized prognostication. This is a case of a 45-year-old male with a right sided convexity meningioma that was resected completely (simpson grade 1). Histopathology demonstrated a meningothelial neoplasm with discohesive cells organized in sheets and a dominance of papillae with a vascular core: WHO Grade III papillary meningioma. Traditionally this patient would be considered high risk for recurrence and would be considered for adjuvant radiation therapy. However, this patient did not receive adjuvant radiation therapy (prolonged complicated post-operative course), and at 62-months after surgical resection the patient remains recurrence-free. The methylation-based predictor score for this patient is 0.9785 (score ranges from 0.004669 [lowest risk] to 1 [highest risk]). (A) The meningioma recurrence score for this patient demonstrates a 19.07% risk of recurrence within 5-years making this patient low risk for recurrence. (B) Copy number variation plot demonstrates gain in chromosome 10 and 22q loss. (C) Existing DNA methylation-classification systems deem this tumor to be of intermediate risk without clear predictions of 5-year recurrence risk.

FIG. 16 shows suitable configured computer device, and associated communications networks, devices, software and firmware to provide a platform for enabling one or more embodiments as described herein.

DETAILED DESCRIPTION

In the following description, numerous specific details are set forth to provide a thorough understanding of the invention. However, it is understood that the invention may be practiced without these specific details.

Variability in standard-of-care classifications preclude accurate predictions of early tumor recurrence for individual patients with meningioma limiting the appropriate selection of patients who would benefit from adjuvant radiotherapy to delay recurrence. We aimed to develop an individualized prediction model of early recurrence risk combining clinical and molecular factors in meningioma.

DNA methylation profiles of clinically-annotated tumor samples across multiple institutions were used to develop a methylome-model of 5-year recurrence-free-survival (RFS). Subsequently, a 5-year meningioma recurrence score was generated using a nomogram that integrated the methylome-model with established prognostic clinical factors. Performance of both models was evaluated and compared to standard-of-care models using multiple independent cohorts.

The methylome-based predictor of 5-year RFS performed favorably compared to a grade-based predictor when tested using the three validation cohorts (ΔAUC=0.10, 95% CI 0.03-0.018) and was independently associated with RFS after adjusting for histopathological grade, extent of resection and burden of copy number alterations (HR 3.6, 95% CI 1.8-7.2, P<0.001). A nomogram combining the methylome-predictor with clinical factors demonstrated greater discrimination than a nomogram using clinical factors alone in two independent validation cohorts (ΔAUC=0.25, 95% CI 0.22-0.27) and resulted in two groups with distinct recurrence patterns (HR 7.7, 95% CI 5.3-11.1, P<0.001) with clinical implications.

The models developed and validated in this study provide important prognostic information not captured by previously established clinical and molecular factors which could be used to individualize decisions regarding post-operative therapeutic interventions, in particular, whether to treat patients with adjuvant radiotherapy versus observation alone.

In an aspect, there is provided a method of predicting recurrence free survival in a patient with meningioma comprising:

(a) determining a tumor DNA methylation profile from a tumor sample from the patient, the tumor DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9;

(b) calculating a risk of meningioma recurrence based on comparing the tumor DNA methylation profile and a reference methylation profile comprising the extent to which the methylation status of the at least 200 loci is associated with a risk of recurrence.

As used herein, “tumour DNA methylation profile” in a given genetic tumour sample is the extent of DNA methylation associated with the tumour DNA. In preferable embodiments, a tumour DNA methylation profile describes the extent of DNA methylation associated with specific loci in tumour DNA

As used herein, “reference methylation profile” is the extent DNA methylation, preferably associated with specific loci, that is associated with a favourable or unfavourable outcomes as they relate to meningioma, for example, the risk of recurrence.

As used herein, “risk of recurrence” means the chance that a cancer that has been treated will recur. Risk recurrence may be stratified risk categories, i.e. low, intermediate, or high.

In some embodiments, the at least 200 loci is at least 500 loci.

In some embodiments, the at least 200 loci is at least 1000 loci.

In some embodiments, the at least 200 loci is at least 1500 loci.

In some embodiments, the at least 200 loci is at least 2000 loci.

In some embodiments, the at least 200 loci is at least 2500 loci.

In some embodiments, the at least 200 loci is at least 3500 loci.

In some embodiments, the at least 200 loci is at least 4000 loci.

In some embodiments, the at least 200 loci is at least 5000 loci.

In some embodiments, the at least 200 loci is at least 6000 loci.

In some embodiments, the at least 200 loci is at least 7000 loci.

In some embodiments, the at least 200 loci is at least 8000 loci.

In some embodiments, the at least 200 loci is at least 9000 loci.

In some embodiments, the at least 200 loci is at least 9500 loci.

In some embodiments, the at least 200 loci is at least 9529 loci.

In some embodiments, calculating the risk is based further on at least one of the WHO grading and Simpson grading of the tumor.

In some embodiments, the method further comprises calculating a predicted number of years of recurrence free survival (RFS Score).

In some embodiments, the method further comprises stratifying the patient into high, medium and low risk categories based on said RFS Score compared to known reference patients.

In some embodiments, the method further comprises treating the patient with adjuvant therapy if the patient is in the high or medium risk category.

In some embodiments, the adjuvant therapy comprises chemotherapy or radiotherapy.

In an aspect, there is provided an array consisting of the 9529 probes set forth in Table 9.

The present system and method may be practiced in various embodiments. A suitably configured computer device, and associated communications networks, devices, software and firmware may provide a platform for enabling one or more embodiments as described above. By way of example, FIG. 16 shows a generic computer device 100 that may include a central processing unit (“CPU”) 102 connected to a storage unit 104 and to a random access memory 106. The CPU 102 may process an operating system 101, application program 103, and data 123. The operating system 101, application program 103, and data 123 may be stored in storage unit 104 and loaded into memory 106, as may be required. Computer device 100 may further include a graphics processing unit (GPU) 122 which is operatively connected to CPU 102 and to memory 106 to offload intensive image processing calculations from CPU 102 and run these calculations in parallel with CPU 102. An operator 107 may interact with the computer device 100 using a video display 108 connected by a video interface 105, and various input/output devices such as a keyboard 115, mouse 112, and disk drive or solid state drive 114 connected by an I/O interface 109. In known manner, the mouse 112 may be configured to control movement of a cursor in the video display 108, and to operate various graphical user interface (GUI) controls appearing in the video display 108 with a mouse button. The disk drive or solid state drive 114 may be configured to accept computer readable media 116. The computer device 100 may form part of a network via a network interface 111, allowing the computer device 100 to communicate with other suitably configured data processing systems (not shown). One or more different types of sensors 135 may be used to receive input from various sources.

The present system and method may be practiced on virtually any manner of computer device including a desktop computer, laptop computer, tablet computer or wireless handheld. The present system and method may also be implemented as a computer-readable/useable medium that includes computer program code to enable one or more computer devices to implement each of the various process steps in a method in accordance with the present invention. In case of more than computer devices performing the entire operation, the computer devices are networked to distribute the various steps of the operation. It is understood that the terms computer-readable medium or computer useable medium comprises one or more of any type of physical embodiment of the program code. In particular, the computer-readable/useable medium can comprise program code embodied on one or more portable storage articles of manufacture (e.g. an optical disc, a magnetic disk, a tape, etc.), on one or more data storage portioned of a computing device, such as memory associated with a computer and/or a storage system.

In an aspect, there is provided a computer program product for use in conjunction with a computer having a processor and a memory connected to the processor, the computer program product comprising a computer readable storage medium having a computer mechanism encoded thereon, wherein the computer program mechanism may be loaded into the memory of the computer and cause the computer to carry out the method described herein.

In an aspect, there is provided a computer implemented product for predicting recurrence free survival in a patient with meningioma comprising:

(a) a means for receiving values corresponding to a tumor DNA methylation profile from a tumor sample from the patient, the DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9;

(b) a database comprising a reference methylation profile representing the extent to which each said methylation status of the at least 200 loci is associated with a risk of recurrence; and (c) a means for calculating a risk of meningioma recurrence based on a comparison of the tumor DNA methylation profile and the reference methylation profile.

In some embodiments, the computer implemented product described herein is for carrying out the method described herein.

In some embodiments, the computer implemented product further outputs a recommendation for adjuvant therapy if the patient has been classified into a medium or high risk group.

In an aspect, there is provided a computer readable medium having stored thereon a data structure for storing the computer implemented product described herein.

In an aspect, there is provided a computer system comprising

(a) a user interface capable of receiving values corresponding to a tumor DNA methylation profile from a tumor sample from the patient, the DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9;

(b) a database including records comprising a reference methylation profile comprising the extent to which each said methylation status of the at least 200 loci is associated with a risk of recurrence;

(c) an output that displays a prediction of recurrence free survival based on a comparison of the tumor DNA methylation profile and the reference methylation profile. In an aspect, there is provided a method of predicting recurrence free survival in a patient with meningioma comprising:

(a) determining a tumor DNA methylation profile from a tumor sample from the patient, the tumor DNA methylation profile comprising the methylation status of at least 200 genes set forth in Table 9;

(b) calculating a risk of meningioma recurrence based on comparing the tumor DNA methylation profile and a reference methylation profile setting forth the extent to which the methylation status of the at least 200 genes is associated with a risk of recurrence.

In some embodiments, the at least 200 genes is at least 400, 600, 800, 1000, 1500, 2000, 2500, 3000, or 3237 genes.

As used herein, “processor” may be any type of processor, such as, for example, any type of general-purpose microprocessor or microcontroller (e.g., an Intel™ x86, PowerPC™, ARM™ processor, or the like), a digital signal processing (DSP) processor, an integrated circuit, a field programmable gate array (FPGA), or any combination thereof.

As used herein “memory” may include a suitable combination of any type of computer memory that is located either internally or externally such as, for example, random-access memory (RAM), read-only memory (ROM), compact disc read-only memory (CDROM), electro-optical memory, magneto-optical memory, erasable programmable read-only memory (EPROM), and electrically-erasable programmable read-only memory (EEPROM), or the like. Portions of memory 102 may be organized using a conventional filesystem, controlled and administered by an operating system governing overall operation of a device.

As used herein, “computer readable storage medium” (also referred to as a machine-readable medium, a processor-readable medium, or a computer usable medium having a computer-readable program code embodied therein) is a medium capable of storing data in a format readable by a computer or machine. The machine-readable medium can be any suitable tangible, non-transitory medium, including magnetic, optical, or electrical storage medium including a diskette, compact disk read only memory (CD-ROM), memory device (volatile or non-volatile), or similar storage mechanism. The computer readable storage medium can contain various sets of instructions, code sequences, configuration information, or other data, which, when executed, cause a processor to perform steps in a method according to an embodiment of the disclosure. Those of ordinary skill in the art will appreciate that other instructions and operations necessary to implement the described implementations can also be stored on the computer readable storage medium. The instructions stored on the computer readable storage medium can be executed by a processor or other suitable processing device, and can interface with circuitry to perform the described tasks.

As used herein, “data structure” a particular way of organizing data in a computer so that it can be used efficiently. Data structures can implement one or more particular abstract data types (ADT), which specify the operations that can be performed on a data structure and the computational complexity of those operations. In comparison, a data structure is a concrete implementation of the specification provided by an ADT.

The advantages of the present invention are further illustrated by the following examples. The examples and their particular details set forth herein are presented for illustration only and should not be construed as a limitation on the claims of the present invention.

EXAMPLES

Methods and Materials

Study Design and Data Sources

This multicenter retrospective study was carried out in accordance with individual institutional ethics and review board guidelines and comprised a total of 486 patients with clinically annotated and available meningioma samples. Institutional waivers of informed consent were obtained due to minimal patient risk associated with this study. Two-hundred and eighty-two (N=282) fresh-frozen or formalin-fixed paraffin embedded (FFPE) meningioma tumor samples from multiple institutions (N=76 from Princess Margaret Cancer Research Centre, Toronto, Canada; N=206 from European centers including University of Heidelberg, Heidelberg, Germany and Goethe-University, Frankfurt, Germany and University of Tubingen, Tubingen, Germany, and University Hospital Zurich, Zurich, Switzerland and Medical University of Vienna, Vienna, Austria) comprised the discovery cohort. which was split into a training cohort (81%, N=228 samples) and first validation cohort (19%, N=54 samples) each balanced for tumor grade, tissue type, recurrence status and time to recurrence. One hundred and forty (N=140) FFPE meningioma tumor samples from a separate institution (MD Anderson Cancer Centre, Houston, USA) were used as a second validation cohort, and sixty-four (N=64) fresh frozen meningioma tumor samples from two other institutions were used as a third validation cohort (N=46 from Princess Margaret Cancer Research Centre; N=18 from The Chinese University of Hong Kong). The sample sets from Europe and MD Anderson comprised the subset of previously published samples for which clinical data (recurrence-free survival [RFS], WHO grade) were available^(11,12). Moreover, TERT promoter mutation status was available on a subset of previously published European samples^(10,11). Gene expression analysis was performed on publicly available microarray data on 98 patients with meningiomas of all grades (GSE16581¹⁴ and GSE9438¹⁵). The outline for the overall study design is demonstrated in FIG. 7.

Definitions

Hematoxylin and eosin (H&E) slides for each patient were reviewed for meningioma diagnosis and WHO grading was performed according to the current WHO 2016 criteria at local institutions by experienced neuropathologists. Tumor recurrence and time to recurrence were the primary outcomes of interest and were collected locally for each sample as previously described^(11,12). Briefly, recurrence was defined as tumor growth following gross-total resection or tumor progression following subtotal resection. Time to recurrence was determined by reviewing post-operative imaging and calculating the duration from the date of surgery to first post-operative imaging documenting tumor recurrence in concordance with documentation in the medical charts. The extent of resection (Simpson grade²) was determined based on the surgeon's operative report in correlation with postoperative cranial imaging.

Generation of an Individualized Methylome-Predictor of 5-Year RFS

For DNA methylation and copy number analysis of the samples, DNA was extracted from each tumor and DNA methylation profiling was performed using Illumina 450k HumanMethylation BeadChip or 850k EPIC arrays as per manufacturer instructions at each institution. Raw data files (*.idat) were imported, processed, and normalized to integrate data from multiple generations of Infinium methylation arrays. Copy number aberrations were inferred from methylation array data¹⁶ and burden of copy number alterations was computed per sample as previously described¹⁷. Probes that were common in both 450k HumanMethylation Beadchip and 850K EPIC arrays were selected as possible features for development of our predictor such that our predictor would be applicable to the landscape of available technologies. We used a multi-step strategy to select the probes to be used in the generation of our predictor (See FIG. 8).

To develop the methylome-based predictor of early meningioma recurrence, we performed generalized boosted regression modelling using the final selected probes in samples from the training cohort to predict 5-year RFS. Boosted regression modelling using WHO grade as a sole feature in the training cohort was also performed and tested in each validation cohort to compare methylation-based predictor performance to a standard-of-care model. Performance of both models was assessed by generating time-dependent receiver operating characteristic (ROC) curves and computing average areas under the ROC curves (AUC) for each validation cohort independently, along with their 95% confidence intervals using the bootstrap resampling method with 10,000 resamples¹⁸.

Methylation probe annotation was performed using the UCSC Genome Browse (GRCh38/hg38 assembly). We used the Functional Annotation Clustering algorithm¹⁹ of Database for Annotation, Visualization, and Integrated Discovery (DAVID)¹⁹ Bioinformatics Resource 6.8 to identify redundant functional clusters represented by genes annotated with a minimum of 5 probes (See FIG. 9). Two publicly available microarray datasets (GSE16581¹⁴ and GSE9438¹⁵) reporting on 22486 genes for 98 patients with meningiomas were pooled to correlate methylation data with gene expression data as an exploratory analysis.

Further details regarding the steps for generation, validation and characterization of the methylome-predictor of 5-year RFS is described below.

Generation of a Meningioma-Recurrence Score

To create a contemporary meningioma recurrence score that could be utilized by clinicians to predict early risk of recurrence for individual patients, we generated a nomogram based on a Cox model that incorporated the methylome-based predictor, WHO grade, and extent of resection using samples from the training cohort and second validation cohort to predict 5-year RFS. Both the training cohort and second validation cohort were used to train this nomogram in order to increase the number of samples available to capture the heterogeneity in the spectrum of data available. It is important to note that as none of the samples used to train this model were used to assess model performance on external validation. The global performance of the meningioma recurrence was assessed by generating time-dependent receiver operating characteristic (ROC) curves and computing average area under the ROC curves (AUC) for the first and third validation cohort independently, along with their 95% confidence intervals using the bootstrap resampling method with 10,000 resamples^(18,20). For comparison, a nomogram that incorporated only WHO grade and extent of resection as the sole features was also developed in similar fashion. Internal validation using bootstrap resampling using 10,000 resamples was also performed. Model calibration was assessed visually by plotting observed event rates against nomogram predicted probabilities for two risk groups. Detailed descriptions on nomogram calculations and use are described below and FIG. 10.

Statistical Analysis

Summary statistics are reported as counts (and proportions) for categorical variables and median (and range) for continuous variables, unless otherwise indicated. Cohort size was determined by availability of samples. Statistical analyses were performed in consultation with two expert biostatisticians (L.P and O.S).

To investigate the clinical relevance of the methylome-based predictor and meningioma recurrence score, distribution of survival times was performed using Kaplan-Meier methods and compared across groups using log-rank testing. The frequency of genome-wide copy number alterations across groups was computed and plotted using a custom algorithm. The performance of a methylome-based predictor was compared to grade-based predictor by computing an average ΔAUC (AUC_(methylome)-AUC_(grade)) and 95% CI from all bootstraps and the performance of the meningioma grade, recurrence score incorporating the methylome predictor was compared to a nomogram excluding the methylome predictor by computing average ΔAUC (AUC_(combined nomogram)-AUC_(clinical only nomogram)) and 95% CI in similar fashion.

Hazard ratios, including 95% confidence intervals (CI), were calculated based on univariable and multivariable Cox regression modeling for the methylome-based predictor and other covariates including WHO grade, extent of resection, and burden of copy number alterations. Proportional hazards assumption was tested by computing Schoenfeld residuals for each covariate and testing was performed according to Grambsch and Therneau²¹ (FIG. 11 and Table 1). Martingale residuals were plotted against methylation predictor probabilities to determine appropriateness of linear modeling (FIG. 12). Sensitivity analyses evaluating the possible confounding effects of TERT promoter mutations, receipt of adjuvant radiation therapy, and center-effects were performed using multivariable Cox regression in samples with available information. Comparison of proportions across groups was completed using χ² test and Fisher Exact test, where appropriate.

Two-sided p-values are reported, and the threshold for statistical significance was set a priori at α=0.05. We used R version 3.3.1 for all statistical analyses, model generation, and model validation.

Methylation Analysis

DNA was extracted from each tumor and DNA methylation profiling was performed using Illumina 450k HumanMethylation BeadChip (Illumina, San Diego, Calif., USA) or 850k EPIC (Illumina, San Diego, Calif., USA) arrays per manufacturer instructions at each institution. Raw data files (*.idat) were imported and processed using R 3.3.1 (The R foundation for Statistical Computing www.R-project.org) minfi package. We performed single-sample Noob (ssNoob) normalization to integrate data from multiple generations of Infinium methylation arrays²⁷. Probes that failed to hybridize (detection p value>0.05) in one or more samples, probes that overlapped with single nucleotide polymorphisms, cross-reactive probes, and probes that localized on X and Y chromosomes were removed from analysis²⁸. Samples with poor quality control were also removed (N=2). Missing values were imputed by boosted trees imputation using Imputation package (https://github.com/jeffwong/imputation). Differentially methylated probes were identified based on post-processed β-values.

Copy Number Analysis

DNA copy number alterations were inferred from DNA methylation data of tumor samples using the Bioconductor package conumee relative to normal cortex DNA methylation data²⁹. Whole genome and chromosome-level plots were generated for each sample and manually inspected by two independent reviewers (F.N, S.S) for copy number alterations. Partial and complete losses or gains of the p and q arm of each chromosome was recorded at a threshold of |0.2|. Discrepancies between reviewers were resolved by consensus, and discussed with a third independent reviewer (M.Y) if necessary. The burden of copy number aberrations per sample was calculated as previously described³⁰. In addition to this, the genome of each sample was split into bins of multiple probes and probe intensity values were combined for copy number annotation. A genome-wide copy number alteration plot demonstrating proportions of gains or losses per bin was produced according to a custom algorithm.

TERT Promoter Sequencing

A subset of samples had panel sequencing by applying a custom hybrid-capture approach (Agilent Technologies, Calif., USA) as previously described³¹.

Machine-Learning Algorithms and Statistical Analyses

Feature Selection

Probes that were common in both 450k HumanMethylation Beadchip and 850K EPIC arrays were selected as initial features for development in our algorithms. Feature selection was performed on samples in the training set only. First, univariable Cox regression analysis for survival using each initial feature was performed to create a rank list of discriminative probes. Probes that were highly correlated with RFS (P-value≤0.01) were selected for further analysis. To mitigate for batch differences in tissue source and center of tissue processing, Principal Component Analysis of the highest 50000, 40000, 30000, and 20000 variables probes of stratified by tissue source and institution in the training cohort (European Fresh frozen tissue, European FFPE tissue, Princess Margaret Cancer Center Fresh frozen tissue) was performed. Technical sources of variation were assessed by computing the F-ratio, and the set of highest variable probes that showed a reduction in the F-ratio for tissue source and tissue type with greatest convergence between the center of gravity and standard deviation was used to determine a common set of discriminatory probes. This approach was used strictly for feature selection without any transformation of data. Further feature refinement was performed using the minimum redundancy maximum relevance ensemble (mRMRe) package (https://cran.r-project.org/web/packages/mRMRe/index.html) that allows for parallelized ensemble features to select a set of relevant, complementary, and reliable features⁶. A minimal importance threshold of |2.5| was used for final feature selection.

The distribution of patient and tumor characteristics in our discovery, first validation, and second validation cohorts are shown in Table 1. Our training dataset contained 228 samples with 341,172 overlapping probes from Illumina 450k HumanMethylation BeadChips and 850k EPIC arrays. Univariate Cox regression for survival using the training set revealed 71538 probes that were highly correlated with RFS (P<0.01). Principal Component Analysis based on the top 30000 variable probes form the 71538 available probes demonstrated the least batch related differences with convergence of centre of gravity and standard deviations between fresh frozen and FFPE samples from the Princess Margaret Cancer Center and samples from the University of Heidelberg (FIG. 7, appendix pp). Of the top 30000 variable probes, 23571 probes were in common in samples from Princess Margaret Cancer Center fresh frozen samples and European FFPE and fresh frozen samples and were selected for further refinement. Parallelized minimum redundancy maximum relevance ensemble revealed a final set of 9529 probes annotated to 3237 known genes that were selected as features for development of a methylome-based predictor of RFS.

Model Generation

To develop the predictor, we performed gradient boosted regression modeling using the final selected probes in samples from the training cohort to create and tune a tumor DNA methylome-based predictor of early (5-year). Gradient boosted modeling is an ensemble technique whereby weak individual tree-based prediction models are combined sequentially over ith number of iterations to produce a strong model for predicting outcome. In brief, at every iteration, a tree is grown, residuals are calculated for the tree, and a regression is run to minimize the residual and weights are updated accordingly. Our implementation is cox-proportion hazard model for right-censored data. Gradient boosted modeling using WHO grade (binarized) as a sole feature in the training cohort was also performed and tested in both validation cohorts for comparison of model performance.

Probe Annotation, Pathway, and Gene Expression Analysis

Probe annotation was performed using the UCSC Genome Browse (GRCh38/hg38 assembly). We used the Functional Annotation Clustering algorithm³³ of Database for Annotation, Visualization, and Integrated Discovery (DAVID)³³ Bioinformatics Resource 6.8 to identify functional clusters represented by genes annotated with a minimum of 5 probes. Functional clustering was performed using high classification stringency. The group enrichment score, which represents the geometric mean of the genes p-values in each cluster, was used to rank biological significance.

To correlate methylation data with gene expression, we pooled 2 publicly available microarray datasets (GSE16581³⁴ and GSE9438³⁵) on 22486 genes for 98 patients with meningiomas. Affymetrix CEL files of each dataset were imported using Partek Genomics Suit Software (Partek, St. Louis, USA) and processed by quantile normalization, log2 transformation, and Robust MultiArray Average background correction. Gene level integration was performed by averaging multiple probes mapped to each gene after inverse log2 transformation for each dataset. Dataset integration was performed after batch effect correction, and fold changes and corresponding p-values of the genes of interest were obtained.

Nomogram Development

We used a Cox model to generate a nomogram combining methylome-based predictor scores, Simpson grade, and WHO grade as variables of interest using samples from the training cohort and validation cohort 2. Given that the development of the model is highly dependent on the input number of samples, we combined the training cohort and validation cohort 2 to generate this model. The tuning parameter λ was trained using 10-fold cross-validation approach with 1 standard error selection rule. The model was externally validated using validation cohort 2 and validation cohort 3 (representing total of 2 rounds validation).

Nomogram Use

A nomogram is a two-dimensional diagram used to allow for approximate graphical calculation of mathematical function using scaled graphical devices. Each variable in the equation is scaled accordingly, and values for each variable are assigned points in the “calculator” (FIG. 9, appendix pp). The variables included into the nomogram are the “5y methylome probability”, “WHO grade” and “Simpson grade”. We are developing a publicly available platform where users can upload their raw IDAT files for their samples to receive the 5-year methylome based predictor probability. For a given patient, the cumulative points from each variable (“5y methylome probability”, “WHO grade” and “Simpson grade”) is calculated for a result of total points. The “risk nomogram” is then used to scale the total points to risk of 5-year RFS. To facilitate unrestricted global dissemination, we have created a freely available online calculator of the meningioma recurrence score (https://fnassiri.shinyapps.io/meningioma_recurrence_score_online_calculator/).

Statistical Analysis

Summary statistics are reported as counts (and proportions) for categorical variables and median (and range) for continuous variables, unless otherwise indicated. Cohort size was determined by availability of samples and not by a priori power calculations. Statistical analyses were performed in consultation with two expert biostatisticians (L.P and O.S).

To investigate the clinical relevance of the 5y methylome-based predictor distribution on survival times, we used Kaplan-Meier methods and comparisons across groups through log-rank testing. Performance of the predictor was assessed by independently computing probability of recurrence at 5 years in each of the 3 validation cohorts and in the combined validation cohorts using 10,000 bootstrap resampling approach. Receiver operator characteristic (ROC) curves were generated for each bootstrap using the Nearest Neighbour time-dependent ROC curve method of Heagerty, Lumley, and Pepe (2000)³⁶ and the area under the ROC curve (AUC) at 5-years was computed for each bootstrap and averaged to calculate the accuracy of our model to predict recurrence at 5 years in all validation cohorts. The performance of a methylome-based predictor was compared to grade-based predictor by computing ΔAUC (AUC_(methylome)-AUC_(grade)) for each bootstrap and averaged to compute a mean ΔAUC with 95% CI. To evaluate the prognostic utility of the methylome predictor in WHO grade I, II and WHO grade III tumors, we generated low and high risk groups by splitting from the median of probability results from the methylome predictor and performing Kaplan-Meier survival analyses.

The performance of the meningioma recurrence score incorporating the methylome predictor was compared to a nomogram excluding the methylome predictor by computing time-dependent AUC and ΔAUC at 5-years in a similar fashion to above^(36,37) Performance of the meningioma recurrence was assessed by independently computing probability of recurrence at 5 years in each of the 2 validation cohorts and in the combined validation cohorts using 10,000 bootstrap resampling approach. Receiver operator characteristic (ROC) curves were generated for each bootstrap and the area under the ROC curve (AUC)³⁷ at 5-years was computed for each bootstrap and averaged to calculate the accuracy of our model to predict recurrence at 5 years in all validation cohorts. The performance of the meningioma recurrence score was compared to a nomogram incorporating clinical factors only by computing ΔAUC (ΔAUC=AUC_(combined nomogram)−AUC_(clinical only nomogram)) for each bootstrap and averaged to compute a mean ΔAUC with 95% CI. Calibration of the meningioma recurrence score was assessed visually by plotting observed event rates against nomogram predicted probabilities for two risk groups (high and low risk). To investigate the clinical relevance of the meningioma-recurrence score, distribution of survival times was estimated using Kaplan-Meier methods and compared across high and low risk groups using log-rank testing.

Hazard ratios and their 95% confidence intervals (CI), were calculated based on univariable and multivariable Cox regression modeling for the methylome-based predictor and other covariates including WHO Grade, extent of resection, and burden of copy number alterations. In order to determine whether TERT promoter mutations. receipt of adjuvant radiation therapy, and treatment center were confounding the relationship between the methylome-based predictor and RFS, sensitivity analyses were reported including these factors as covariates where available. Plots of martingale residuals versus methylome-predictor values were generated to assess the appropriateness of the assumption of linear effect of the methylome-based predictor in the Cox regression model. The proportional hazards assumptions were evaluated by computing plots of Schoenfeld residuals for each covariate versus time. Weighted least-squares line were fitted to the residual plot, and proportional hazards testing was performed according to Grambsch and Therneau¹².

Results and Discussion

A set of 9529 probes were selected from an initial training cohort and generalized boosted regression modeling was performed to develop a DNA methylation-based predictor of 5-year recurrence risk in meningioma (FIG. 7). Three validation cohorts (Table 2) were used to test the performance of the methylation-based predictor compared to a grade-based predictor.

Validation of a Methylome-Predictor of Early Meningioma Recurrence

The methylome-based predictor performed favorably compared to the grade-based predictor at 5-years in each of the 3 validation cohorts (FIG. 1) with substantial statistical improvements in performance when tested in all combined validation cohorts (ΔAUC=0.10, 95% CI 0.03-0.18). When stratified by median, the 5-year methylome-predictor distinguished risk groups (lower and higher risk) in all 3 validation cohorts (FIG. 2). Patients in the higher risk group had a median RFS of 2.1 years, 8.1 years, and 4.2 years in the first, second, and third validation cohorts respectively compared to patients in the lower risk groups which had median RFS of “unreached” in the first and second validation cohorts and median RFS of 7.2 years in the third validation cohort (HR 8.1 95% CI 2.8-23.8, HR 3.5 95% CI 1.8-6.6, and HR 2.0, 95% CI 1.2-3.7, respectively).

DNA copy number analysis demonstrated that increased frequency of copy number aberrations in the higher risk groups, as demonstrated by high proportion of chromosomal deletions in 1p, 4p, 6q, 10q, 14q, and 18q (FIG. 3). The total burden of copy number alterations was also correlated with risk groups, with greatest proportion of burden of copy number alterations found in the higher risk groups in all three validation cohorts (FIG. 9). It is noteworthy that of all patients with high burden of copy number aberrations from all three validation cohorts, only 12 patients (4.6%) were in the lower risk group.

Multivariable Cox regression analysis demonstrated that the 5-year methylome based predictor was independently associated with RFS in samples from all validation cohorts (HR 3.6, 95% CI 1.8-7.2, P<0.001) after controlling for tumor grade, extent of resection, burden of copy number alterations (Table S4). Sensitivity analyses including receipt of adjuvant therapy, TERT promoter mutations, and center of treatment as covariates did not alter this relationship and these covariates were not independently associated with RFS (Tables 5-7).

Current clinical practice relies on histopathologic grade to inform the decision of post-surgical management of meningioma. Patients with grade I tumors are most commonly monitored with serial imaging after surgery, while patients with grade II and III tumors are considered for adjuvant therapy, such as radiation, to prevent recurrence. There is, however, unexplained clinical variability in RFS within all grades of meningioma, and we examined the use of the 5-year methylome-predictor to address this issue. Among patients with WHO grade II tumors in all validation cohorts, the median RFS for the higher risk group was 2.6 years compared to a median RFS of 8.4 years in the lower risk group (HR 2.8, 95% CI 1.7-4.8, P<0.001, FIG. 4B). Patients with WHO Grade I tumors in all validation cohorts also had increased risk for recurrence in the higher risk group (HR 2.9, 95% CI 1.3-6.6, P=0.006, FIG. 4A) with only 3 patients in the lower risk group recurring within the first 5 years compared to 18 patients in the higher risk group. Lastly, patients with WHO Grade III tumors from all validation cohorts had poor median RFS in the higher risk group (1.3 years), compared to a median RFS of 6.0 years in the lower risk group (HR 3.0, 95% CI 1.4-6.5, P=0.004, FIG. 4C).

Characterization of Predictor CpG Sites

The selected 9529 probes used in our model comprise only 1% of all probes included on the 850K Illumina Array. These probes were enriched to be found in the promoter regions (N=3057, 32.1%) and located on CpG islands (N=4633, 48.6%) compared to all probes found on the 850K Illumina Array (29.6%, P<0.001 and 18.0%, P<0.001, respectively; Table 8) Of the 9529 probes, only 1261 (13.2%) were “favorable” probes associated with lower risk of recurrence when methylated (HR ranging from 0.002 to 0.38, on univariable Cox regression with associated P<0.001). The remaining 8237 (86.8%) probes were “unfavorable” probes associated with higher risk of recurrence when methylated (HR ranging from 2.45 to 517.82, on univariable Cox regression with associated P<0.001).

There were 2332 probes annotated to 294 genes with at least 5 probes represented per gene. Of these (Table 9), only 68 of 2332 probes (2.9%) were found to be “favorable” probes (associated with lower recurrence risk when methylated, HR ranging from 0.008 to 0.252 on univariable Cox regression with associated P<0.001). The remaining 2265 probes (97.1%) were found to be “unfavorable” probes associated with greater recurrence risk (associated with higher recurrence risk when methylated, HR ranging from 2.95 to 517.82 on univariable Cox regression with associated P<0.001). Functional annotation clustering of these 2265 “unfavorable” probes revealed that homeobox (enrichment score=54.33) and TBox (enrichment score=4.12) were highly significant redundant functional clusters (FIG. 9). Gene expression analysis of Homeobox family of genes and Tbox genes for which methylation data was also available, demonstrated that although these genes were relatively hypermethylated in recurrence-prone tumors, they were either upregulated or non-differentially expressed (Table 10).

Validation and Clinical Utility of a Meningioma Recurrence Score

To generate and validate a meningioma recurrence score that could be translated to the clinic, we developed a nomogram to predict 5-year RFS that incorporated the validated 5y methylome-based predictor with established prognostic covariates (WHO grade, and extent of resection) using samples from the training cohort and second validation cohort (FIG. 5). Validation of the nomogram using the first validation cohort and third validation cohort independently as well as both validation cohorts combined demonstrated greater predictive performance of the nomogram with methylome predictor included compared to a nomogram using clinical factors (WHO grade and extent of resection) alone (ΔAUC=0.24, 95% CI 0.19-0.28; ΔAUC=0.26, 95% CI 0.22-0.31; ΔAUC=0.25, 95% CI 0.22-0.27 respectively, FIG. 5). The discriminative power of the meningioma recurrence score was approximately 82% in combined validation cohorts (AUC=0.82, 95% CI 0.76-0.87). The performance of the model using external validation and internal validation approaches were similar (FIG. 13). The highest proportion of allottable points given in the nomogram is based on probabilities from the methylome-based predictor, again suggesting that the methylome-predictor has greater importance in determining recurrence risk in meningiomas compared to established clinical factors. Calibration and Kaplan-Meier survival analysis of the meningioma recurrence score clearly stratifies patients with high risk and low risk for 5-year recurrence (HR 7.7, 95% CI 5.3-11.1, P<0.001, FIG. 6). Interestingly, while histologic grades II and III are meant to predict high recurrence risk, we find that the low-risk group in total contains 39 grade II tumors (34.5% of the grade II tumors) and 4 grade III tumors (8.3% of the grade III tumors).

Conversely, while a WHO grade I designation is meant to convey a low risk of recurrence, there were 35 (21.2%) patients with grade I tumors in the high-risk group. These results indicate refinement of risk estimate by the nomogram relative to current classification standards. To facilitate unrestricted global dissemination, we have created a freely available online calculator of the meningioma recurrence score https://meningiomas.shinyappsio/meningioma_recurrence_score_online_calculator/.

Details on imputations for the online calculator can be found in the methods above, and case examples demonstrating the power of personalized predictions are detailed in FIGS. 14-16.

Discussion

In this multicenter study, we demonstrated the transformative utility of integrating clinical and molecular factors for use beyond simple classification into the realm of individualized prognostication in neuro-oncology. Our methylome-based predictor was more reliably able to predict early (5-year) recurrence-free survival in comparison with histologic grading and was associated with RFS independent of established clinical and molecular factors. Combining the methylome predictor with established prognostic clinical factors (WHO grade and extent of resection) in a meningioma recurrence score refined prognostication for individual patients with meningiomas beyond established prognostic clinical and molecular factors with therapeutic implications for individualizing decision-making regarding the need for adjuvant therapy after surgery in meningiomas.

Although WHO grade is associated with recurrence in populations of meningioma patients, and is currently used to guide therapy, the clear within-grade variation for risk of recurrence and inter-rater variability makes it challenging to rely on tumor grade alone to predict recurrence and guide postoperative management decisions for individual patients. As a manifestation of this imprecision, some patients with biologically aggressive tumors may be inappropriately subsumed in within the group of histologically benign tumors. With current standard of care, it is thought that up-front treatment with adjuvant radiation therapy after surgical resection offers the best chance to delay recurrence, and therefore some patients are not appropriately selected for adjuvant treatment with standard-of-care approaches^(3,4). Conversely, there are also some patients with histologically defined intermediate or higher-grade tumors that in fact harbor indolent tumor behavior. Such patients may be receiving adjuvant radiation therapy in the absence of a defined need. Radiation, even when optimized to minimize adverse effects, still carries the risk for adverse radiation effects such as reactive inflammation, vasculitis, and necrosis, all of which have sequelae on patient cognition and quality of life⁵. There is a clear need for a more refined predictor of recurrence patterns for individual patients with meningiomas beyond simple classifications, so that the decision for adjuvant therapy, radiotherapy or otherwise, can be appropriately selected and personalized for patients.

The burden of chromosomal alterations have repeatedly been shown to be one of the most important prognostic molecular alterations in meningioma^(11,17,22.) Similar to others^(11,22), we found recurrent alterations in 1p, 4p, 6q, 10q, 14q, and 18q in a subset of meningiomas that were enriched in higher risk group. However, using a previously published copy number score designed to identify meningiomas with high recurrence risk¹⁷, we found that the burden of copy number alteration was not an independent predictor of recurrence once adjusted for methylation signature. Similarly, although TERT promoter mutations are known to be enriched in more aggressive meningiomas¹⁰, our analysis demonstrated that TERT promoter mutation was not independently associated with RFS on multivariable analysis with the 5-year methylome predictor included in the model. Taken together, these results suggest that our 5-year methylome predictor can provide prognostic information beyond previously established molecular factors in meningioma.

Probes included in the predictor were selected based on correlations of either hypo- or hypermethylated status with respect to RFS. Interestingly, the distribution of these probes was not random, either with respect to methylation status nor with respect to association with known genes. The majority (over 86%) of our included probes were all associated with unfavorable RFS when hypermethylated. This suggests that in general, hypermethylation of a set of CpG sites in meningioma correlates with clinical aggressiveness. For example, hypermethylation of the homeobox and Tbox families of genes were found to be highly overrepresented in the set of relevant probes, suggesting possible involvement of this class of developmental factors in the clinical behavior of meningioma, which requires further validation.

To generate a tool that could be used by clinicians to capture the heterogeneity in recurrence risk in meningiomas, we established a 5-year meningioma recurrence score that combined our validated methylome-based predictor with well-established prognostic clinical factors (WHO grade and extent of resection). The performance of our meningioma recurrence score was improved with the methylome-based predictor included in the nomogram (ΔAUC=0.25, 95% CI 0.22-0.27), and the overall discrimination of our nomogram was high (AUC approximately 82% in two independent cohorts). The construction, and evaluation, of our nomogram meets the standards of reporting on nomograms in oncology and is one of few to demonstrate robust evaluation using multiple independent validation cohorts²³. Now that we have demonstrated that our tools are robustly validated, we are well positioned to prospectively validate the use of our tools to demonstrate efficacy with adjuvant therapy strategies in high-risk patients²⁴. Our meningioma recurrence score informs both patients and clinicians about individualized risk of recurrence, can be used to guide clinicians regarding the need for adjuvant therapy and/or close clinical follow-up.

Our study has some limitations. First, although we have identified a group of probes with distinct epigenetic changes that in combination are predictive of recurrence risk in meningiomas for individual patients, it is unclear whether these changes may be conferring variant behavioural phenotypes or whether they are a surrogate for general cellular dysregulation. Nevertheless, the set of highly refined and selected probes in our predictor are enriched to be located on promotors of CpG islands where aberrant DNA methylation has clearly been linked to carcinogenesis²⁵. The correlation of methylation with gene expression in our study was exploratory and would benefit from additional investigation with matched epigenetic and transcriptomic analysis in the same samples. Moreover, although each institution conformed to a common definition for tumour recurrence and time to recurrence, there is no universally standardized definition of recurrence in meningiomas. The Response Assessment in Neuro-Oncology guidelines may offer an avenue for standardized definition of recurrence to be collected on meningiomas in future clinical trials, which will help with communication across different centres and may also help with further model refinement²⁶.

Our predictor has been designed and validated such that it can be applied to data from fresh frozen or paraffin embedded tissues using the current commonly-used standard platform for genome-wide DNA methylation profiling, facilitating immediate adoption into clinical practice. Our newly developed and retrospectively validated meningioma recurrence score combines both methylome and clinical factors and can be freely used by clinicians to personalize decision making regarding post-operative management of the most common primary intracranial brain tumor via a web-based interface.

TABLE 1 Grambsch and Therneau Proportional Hazards Testing for Cox modeling Covariate Rho Chi-Sq P-value Grade = G2 0.1376 1.807 0.1789 Grade = G3 0.0855 0.669 0.4134 Simpson = Subtotal 0.0832 0.649 0.4205 Meth_predict_probability −0.2036 3.184 0.0744 CNV_Load = Low −0.1574 2.593 0.1073 GLOBAL NA 5.787 0.3275

TABLE 2 Distribution of patient and tumor characteristics used for training and validation. First Validation Second Validation Third Validation Training Cohort cohort Cohort Cohort (n = 228) (n = 54) (n = 140) (n = 64) Source Princess 62 (27%) 14 (26%) 0 (0%) 46 (72%) Margaret Cancer Centre European 166 (73%) 40 (74%) 0 (0%) 0 Centres M D Anderson 0 (0%) 0 (0%) 140 (100%) 0 Cancer Centre Chinese 0 0 0 18 (28%) University of Hong Kong Tissue Type FFPE 118 (52%) 31 (57%) 140 (100%) 0 (0%) Fresh Frozen 110 (48%) 23 (43%) 0 (0%) 64 (100%) WHO Grade WHO I 103 (45%) 24 (44%) 90 (64%) 14 (22%) WHO II 91 (40%) 22 (41%) 43 (31%) 32 (50%) WHO III 34 (15%) 8 (15%) 7 (5%) 18 (28%) Simpson Gradeϕ 1 86 (43%) 18 (43%) 69 (53%) 8 (15%) 2 71 (36%) 14 (33%) 36 (28%) 18 (25%) 3 2 (11%) 4 (10%) 9 (7%) 6 (12%) 4 19 (10%) 6 (14%) 15 (12%) 19 (37%) 5 1 (1%) 0 (0%) 0 (0%) 0 (0%) Recurrence Yes 92 (40%) 21 (39%) 93 (66%) 50 (78%) No 136 (60%) 33 (61%) 47 (34%) 14 (22%) Time to 4.8 (0.1-52) 6.2 (0.25-22.9) 7.0 (0.1-17.6) 5.3 (0.1-18.5) recurrence or last follow-up Genderφ Female 111 (66%) 23 (62%) 98 (70%) 34 (53%) Male 57 (34%) 14 (38%) 42 (30%) 30 (47%) ϕProportions are calculated based on available data (29 missing from training cohort, 12 missing from first validation cohort, 11 missing from second validation cohort, 13 missing from third validation cohort) φProportions are calculated based on available data (60 missing from training cohort, 17 missing from first validation cohort)

TABLE 3 Distribution of burden of copy number alterations among risk groups defined by the 5-year methylome-based predictor. High Low burden of burden of Risk copy number copy number P- Cohort Group alterations alterations Value First Low 2 25 <0.0001 Validation risk Cohort High 16 11 risk Second Low 3 67 0.0099 Validation risk Cohort High 32 38 risk Third Low 7 25 0.0097 Validation risk Cohort High 18 14 risk

TABLE 4 Results of univariable and multivariable cox proportional hazards model for 5 y methylation-based RFS predictor and associated covariates for all validation cohorts. Univariable Cox Multivariable Cox Covariate HR (95% CI) P HR (95% CI) P 5 y methylation- 8.5 (5.3-13.5) <0.001 3.6 (1.8-7.2) <0.001 based predictor WHO Grade <0.001 Grade II vs I 4.0 (2.6-6.2) <0.001 2.6 (1.6-4.4) Grade III vs I 7.1 (4.2-11.8) <0.001 3.1 (1.6-6.2) Grade III vs II 1.8 (1.1-2.7) 0.010 1.2 (0.7-2.0) Extent of Resection 2.2 (1.4-3.5) <0.001 1.6 (1.0-2.5) 0.050 (subtotal vs gross total) burden of copy 2.9 (2.0-4.1) <0.001 1.3 (0.8-2.1) 0.36 number alterations (high vs low) HR, Hazard Ratio; CI, Confidence Interval; P, P-value; WHO, World Health Organization

TABLE 5 Multivariable cox regression analysis including receipt of adjuvant radiation therapy as a covariate the third validation cohort. Multivariable Cox Covariates HR lower .95 upper .95 P Binary Methylome 3.6 1.2 11.0 0.02 Classifier WHO Grade II vs I 2.4 0.8 6.7 0.10 WHO Grade III vs I 1.6 0.5 5.5 0.43 Extent of Resection 1.0 0.5 2.2 0.93 (subtotal vs gross total) Copy number burden 0.8 0.4 1.7 0.57 (high vs low) Adjuvant radiotherapy 1.1 0.3 5.3 0.82 (no vs yes)

TABLE 6 Univariate and multivariable cox regression analysis including TERT promoter mutation status as a covariate in 103 samples. Univariate Cox Multivariate Cox Covariates HR (95% CI) P HR (95% CI) P Binary Methylome 49.6 (17.3-142.3) <0.001 40.2 (11.8-136.4) <0.001 Classifier WHO Grade II vs I 10.3 (3.6-29) <0.001 3.7 (1.2-12.1) 0.02 WHO Grade III vs I 22.7 (7.4-69.8) <0.001 3.1 (0.8-11.3) 0.02 Extent of Resection 2.1 (0.95-4.7) 0.065 2.6 (1.0-6.4) 0.05 (subtotal vs gross total) Copy number burden 4 (2.2-7.3) <0.001 0.8 (0.4-1.6) 0.58 (high vs low) Tert_Mutation (yes vs no) 2.3 (0.56-9.5) 0.25 1.7 (0.3-8.2) 0.54 Gender (male vs female) 1.3 (0.57-3) 0.52 NA NA

TABLE 7 Multivariable cox regression analysis including center of treatment as covariate. Covariate HR (95% CI) PVal 5 y Methylome Predictor 3.15 (1.5-6.5) 0.001 Grade Grade 2 vs1 2.64 (1.5-4.5) 0.001 Grade 3 vs 1 3.3 (1.5-7.2) Extent of resection (subtotal vs gross 1.38 (0.8-2.4) 0.233 total) CNV load (high vs low) 1.32 (0.8-2.2) 0.289 Center Toronto vs Houston 1.16 (0.67-1.99) 0.736 Europe vs Houston 1.43 (0.50-4.07) Hong Kong vs Houston 0.76 (0.28-2.03)

TABLE 8 Annotation of the location of 9529 probes selected in the methylome-predictor in comparison to all probes on 850K EPIC array relative to gene region and CpG island location 9529 probes of 867867 probes methylome on 850K EPIC Annotation predictor array P-value Relative Body 3018 (31.7%) 315159 (36.3%) <0.001 to Gene Unknown 2470 (25.9%) 233769 (26.9%) 0.02 Region Promoter 200-1500 BP 1537 (16.1%) 110079 (13.5%) <0.001 upstream TSS 5′ Untranslated 774 (8.1%) 77468 (8.9%) 0.006 Region 0-200 BP upstream 746 (7.8%) 62439 (7.2%) 0.017 TSS 1^(st) Exon 643 (6.7%) 28604 (3.3%) <0.01 3′ Untranslated Region 247 (2.7%) 22518 (2.6%) 0.987 Exon Band 94 (1.0%) 10831 (1.2%) 0.022 Relative Island 4633 (48.6%) 155862 (18.0%) <0.001 to CpG Not annotated 1470 (15.4%) 488877 (56.3%) <0.001 Island South Shore 1318 (13.8%) 71784 (8.3%) <0.001 North Shore 1172 (12.3%) 74446 (8.6%) <0.001 South Shelf 471 (4.9%) 38500 (4.4%) 0.016 North Shelf 465 (4.9%) 38398 (4.4%) 0.031 Bp, base-pairs; TSS = Transcription Start Site

TABLE 9 Relation to UCSC UCSC Average Average CpG RefGene Univariate P-value Bvalue Bvalue Source Gene Symbol Island Group COX HR COX NotRecurrent Recurrent cg00015770 ORFPR Island 1stExon 14.3473 9.95E−09 0.133082675 0.281372515 cg00018128 TBR1 S_Shelf Body 11.0144 4.04E−10 0.238240155 0.420623714 cg00025981 RGCC Island TSS1500 13.4779 2.04E−07 0.278465395 0.400282216 cg00028678 ARHGEF4 S_Shore TSS1500 7.83729 1.72E−06 0.374075353 0.527289047 cg00035316 HOXD8 Island TSS1500 8.19499 1.75E−07 0.259222373 0.402969469 cg00036347 C2CD4D S_Shore 5URT 6.32633 0.000335 0.529091437 0.640249799 cg00048759 GPC2 Island TSS1500 22.4346 5.62E−07 0.500048411 0.59538254 cg00055679 RNF219-AS1 Island Body 7.3267 0.001505 0.263971124 0.319811879 cg00060304 DKFZp586K1684 Island TSS1500 6.85957 3.35E−07 0.145730157 0.279207375 cg00063471 TNFAIP8L3 Island Body 4.61555 9.76E−06 0.173224235 0.322641626 cg00073003 SOX1 N_Shore 3UTR 4.47482 0.001662 0.179921581 0.246076945 cg00099399 PTPRN2 Island Body 5.48057 0.00105 0.128069916 0.192229567 cg00117172 RUNX3 Island Body 3.27872 0.004437 0.208241782 0.307430286 cg00123473 ZNF518B Island 5URT 7.47258 0.000952 0.598658535 0.682374916 cg00130808 LOC145845 N_Shore TSS200 6.13389 3.35E−05 0.550275743 0.667232657 cg00143249 MNX1 Island Body 11.927 1.68E−07 0.120366053 0.221959892 cg00155423 WNT10A S_Shore 3UTR 4.56435 1.20E−05 0.247210136 0.400766035 cg00155526 NPR3 Island Body 9.51657 1.13E−10 0.128321035 0.317251422 cg00158122 LINC01475 Island Body 8.44283 1.93E−05 0.211559824 0.290543888 cg00168835 BATF3 S_Shore TSS1500 10.9964 1.44E−08 0.157700387 0.327996862 cg00182639 TBX5 S_Shore 5URT 9.07991 3.62E−05 0.267472135 0.358273393 cg00184452 DMRTA2 Island 5URT 9.26715 9.20E−06 0.369912535 0.488499566 cg00204465 MYO15B Island TSS200 18.2843 1.65E−07 0.432291532 0.569108097 cg00204782 TBX1 Island Body 7.11802 0.000169 0.277246944 0.352428077 cg00222472 PBX1 Island Body 53.3916 7.01E−10 0.626374364 0.779958919 cg00237475 SKOR1 Island Body 41.9155 1.70E−11 0.286564279 0.402228939 cg00247571 DLEU1 S_Shelf Body 12.6755 9.44E−06 0.619350287 0.736634889 cg00254681 PRRT1 Island Body 4.91582 0.000384 0.45187213 0.574644258 cg00255368 ESRRG Island 5URT 24.9928 1.04E−09 0.201514705 0.301703141 cg00262621 MMP23B Island TSS1500 20.2182 2.11E−11 0.322781532 0.514607172 cg00286125 HSPB6 Island Body 7.51548 0.0061 0.2262581 0.307019648 cg00298065 ZNF274 Island 5URT 19.2971 2.29E−06 0.441851876 0.527302697 cg00299047 AMH Island 3UTR 8.71985 5.28E−06 0.363207318 0.482147364 cg00299972 TLX1 N_Shelf Body 5.5743 0.000114 0.222249019 0.311564492 cg00306911 FAM184B S_Shore TSS1500 15.1553 4.00E−07 0.467489528 0.608092889 cg00323305 THRB-AS1 Island Body 5.66558 0.000772 0.306803767 0.410433175 cg00357466 AOAMB S_Shore TSS200 9.14915 0.001243 0.483277667 0.566343525 cg00360794 LTK Island Body 6.26784 9.08E−05 0.231788209 0.325944037 cg00384539 PRDM14 Island TSS200 9.99496 7.02E−08 0.279072411 0.449507265 cg00407545 GATA3-AS1 Island Body 3.46913 0.000579 0.317624764 0.442800135 cg00418216 HOXA11-AS N_Shelf Body 10.8103 1.89E−07 0.24663533 0.362670161 cg00425918 NKX2-6 Island Body 9.38265 5.96E−07 0.204725219 0.31011622 cg00431187 HOXAB N_Shore 5URT 6.00991 0.000107 0.542800519 0.649365232 cg00434010 SIM1 Island Body 8.48419 0.003568 0.305682189 0.394276621 cg00435063 PTPRN2 Island Body 5.33199 0.001217 0.587097132 0.655655859 cg00445405 OTX1 S_Shelf Body 8.33913 0.000845 0.61314032 0.699363943 cg00463767 OTX1 N_Shelf Body 13.8807 9.82E−07 0.386075581 0.510870212 cg00487737 ACP5 Island 5URT 7.60446 8.30E−05 0.283832876 0.389328057 cg00490421 CPM N_Shore Body 8.80887 2.79E−05 0.121409633 0.197785945 cg00495775 HOXD11 Island TSS200 29.6619 4.66E−09 0.361394388 0.482595971 cg00499700 F2RL1 Island Body 34.626 7.96E−07 0.580510938 0.702986657 cg00503840 DLX5 S_Shelf Body 10.5934 8.43E−06 0.36137724 0.455306939 cg00504902 HLA-F S_Shore Body 38.2707 3.47E−07 0.618025124 0.732500929 cg00505979 BATF8 Island TSS1500 18.1933 4.36E−09 0.190875209 0.332468709 cg00506343 HOXC6 N_Shelf TSS1500 4.73712 0.002987 0.187823819 0.255529205 cg00507727 ICA1 Island TSS1500 6.64856 1.05E−06 0.132393885 0.281706708 cg00513205 NEtJROG1 S_Shore TSS1500 5.27112 0.000329 0.39426914 0.490526892 cg00553149 GPC2 Island TSS1500 18.1344 9.34E−09 0.212716633 0.338965681 cg00556112 DRD4 Island TSS200 7.85766 1.58E−06 0.105448181 0.21671902 cg00567703 HOXC6 Island 5URT 11.0544 0.001377 0.651858618 0.695643606 cg00574563 LHX1 Island Body 13.5746 2.14E−06 0.178375121 0.279195108 cg00585714 GDF6 Island Body 22.2416 1.59E−07 0.884586736 0.485336515 cg00599564 HCG9 N_Shore Body 9.96976 3.16E−06 0.315296405 0.455941313 cg00616687 SIM2 Island 1stExon 8.69529 1.99E−06 0.13890162 0.250153308 cg00629427 LOC101927248 Island Body 7.72239 0.0027 0.256282319 0.315180947 cg00663077 ZFP42 Island TSS1500 18.3122 1.98E−11 0.255874873 0.442182767 cg00667789 TACSTD2 Island 1stExon 8.87345 9.19E−09 0.15310286 0.314582388 cg00673290 APOB Island Body 5.25098 0.000244 0.223405913 0.332389431 cg00674304 FOMC Island TSS200 5.27037 6.75E−07 0.168430801 0.3556692 cg00687252 IER3 Island Body 15.5542 1.52E−08 0.292954449 0.478726894 cg00714725 CYP26C1 Island Body 5.42767 4.62E−06 0.321032609 0.4883889 cg00716277 PTPRN2 Island Body 12.9091 0.000588 0.627002504 0.678445412 cg00733728 RNF219-AS1 Island Body 12.7367 1.43E−09 0.147365766 0.281130976 cg00756451 TBX5 N_Shelf 5URT 4.87224 0.001593 0.116043461 0.183096098 cg00762160 PAX9 S_Shore TSS1500 4.68286 0.000734 0.292460912 0.394197498 cg00767496 GATM S_Shore TSS1500 4.70187 0.002308 0.528059922 0.624643222 cg00779924 GATA3-AS1 Island Body 3.25869 0.000528 0.259543895 0.403046957 cg00790395 CTP26C1 Island Body 3.42408 0.009074 0.141589595 0.206579638 cg00795248 LHX9 S_Shore Body 6.53516 0.001004 0.142651392 0.204692423 cg00806704 CDX2 N_Shore Body 6.69678 4.09E−06 0.226390116 0.353402851 cg00838919 LHX4 Island Body 4.37342 0.000467 0.210484062 0.311900389 cg00840332 LEP Island TSS200 19.4166 3.46E−08 0.110332463 0.199627364 cg00864474 NTN1 Island 3UTR 8.22807 4.07E−05 0.416496222 0.553032606 cg00867835 SSPO N_Shelf Body 4.84487 0.002995 0.520465853 0.59548602 cg00877329 HPSE2 S_Shelf TSS1500 3.80475 0.000822 0.390047555 0.508399569 cg00882235 LHX4 Island Body 4.1817 0.000724 0.174457011 0.275198746 cg00887814 MGAT4D Island Body 9.51895 3.00E−05 0.219418518 0.308325924 cg00905524 HOXA10-HOXA9 N_Shore Body 17.5414 1.01E−06 0.23800934 0.334874145 cg00913949 ESRRG Island TSS1500 6.09855 3.51E−07 0.157221884 0.309361008 cg00926215 PAX3 S_Shore Body 9.13898 1.99E−07 0.249208929 0.366095146 cg00928397 SP9 N_Shore TSS1500 12.9978 2.69E−06 0.292032191 0.386723358 cg00947782 RNF39 Island Body 5.61442 1.19E−05 0.182835554 0.304016971 cg00949753 PROX1-AS1 N_Shelf Body 7.39327 6.34E−05 0.306086752 0.41201897 cg00963169 ELAVL4 Island Body 8.81412 6.00E−08 0.146744189 0.276138886 cg00970313 PAX9 S_Shore TSS1500 5.5904 2.00E−06 0.210648826 0.372550414 cg00982919 LHX2 Island Body 6.85613 7.27E−05 0.242806392 0.340290604 cg00982984 SHF S_Shelf TSS200 40.0921 2.70E−07 0.518320116 0.649603374 cg00983437 TCF15 S_Shore TSS1500 20.8662 4.04E−07 0.253828081 0.37306729 cg00985729 IER3 S_Shore TSS1500 9.63816 1.55E−05 0.444252713 0.586434717 cg00989858 IRF6 Island 1stExon 11.0236 1.18E−11 0.244929281 0.480542075 cg01003803 FGF2O S_Shore 1stExon 7.92263 9.04E−07 0.163873199 0.289283328 cg01031400 TNFRSF10D Island 1stExon 15.099 2.48E−07 0.353605187 0.504678649 cg01040654 NKX2-5 S_Shelf TSS1500 19.4478 1.52E−06 0.475049721 0.598085697 cg01054755 PROB1 Island 1stExon 4.64664 3.28E−05 0.185086912 0.394057883 cg01078824 HOXA10-HOXA9 S_Shore TSS200 23.8284 7.79E−07 0.388179169 0.474411404 cg01141459 HORMAD2 Island 1stExon 12.3408 5.29E−11 0.289014186 0.475438706 cg01156550 ASCL2 Island TSS1500 7.62818 0.000133 0.321988195 0.415341281 cg01163842 GSC Island Body 6.00845 4.52E−08 0.16968596 0.357688175 cg01172183 STK3 Island 5URT 6.86487 0.000136 0.210523216 0.297804929 cg01174140 SCG33A1 Island TSS1500 61.2054 2.63E−09 0.38314558 0.51960802 cg01177854 ESRP2 Island TSS1500 7.29629 0.002198 0.340034271 0.403061717 cg01182865 GATA4 S_Shore 5URT 4.84361 0.005784 0.3635323 0.430418091 cg01229452 CHRN31 S_Shore Body 12.3569 0.00012 0.332403204 0.429466364 cg01243879 PRR18 Island TSS1500 4.35388 0.004791 0.556669002 0.644357909 cg01258751 SIM1 N_Shelf TSS1500 8.06674 4.45E−06 0.18555699 0.225022553 cg01277331 WNT10A Island Body 42.1238 8.38E−08 0.462205953 0.585142222 cg01277372 MECOM N_Shore 5URT 5.07848 0.001534 0.219323731 0.295871049 cg01285501 HOXA9 Island TSS1500 11.8904 8.64E−06 0.258052078 0.354405145 cg01286685 RNF39 N_Shelf Body 9.47702 4.65E−06 0.53858096 0.338791707 cg01287476 M1AP Island TSS1500 5.89148 0.001394 0.493774202 0.583026211 cg01295203 PRDM14 Island TSS1500 8.30724 2.01E−06 0.296310646 0.422670067 cg01323777 KCNA33 N_Shelf TSS200 5.12493 0.007113 0.762060211 0.835849448 cg01351315 HOXB-AS3 N_Shore TSS200 9.11815 0.000149 0.614287039 0.713299303 cg01371072 PITX2 S_Shelf Body 8.89022 0.000324 0.283985433 0.34386531 cg01402735 ADGRL4 S_Shore TSS200 4.39356 0.009244 0.305126542 0.377621437 cg01405040 HOXD12 S_Shore TSS1500 8.74668 2.36E−08 0.201324534 0.359307049 cg01405445 HSPB6 Island 3UTR 6.67282 1.03E−05 0.503580605 0.666384496 cg01438365 TBX2 Island Body 7.73761 2.50E−05 0.512092655 0.631568949 cg01449663 NTN1 Island Body 77.0292 6.42E−14 0.308803 0.488207879 cg01456368 DNAH1 5URT 13.6129 7.24E−08 0.166824068 0.283797049 cg01460436 SLC12A5 Island 3UTR 19.6905 1.16E−09 0.3067875 0.456929925 cg01463565 NTN1 Island Body 15.8196 1.03E−10 0.089877795 0.227808991 cg01467266 JSRP1 Island Body 7.80021 6.22E−06 0.363951202 0.491581073 cg01484487 LBX2 Island Body 6.05421 0.000145 0.230136183 0.335505836 cg01494454 BCAT1 N_Shore 1stExon 14.433 7.85E−08 0.202156688 0.319796983 cg01499197 CDX2 Island TSS1500 7.75673 5.51E−05 0.261030774 0.348812792 cg01530101 KCNQ1DN Island TSS200 6.74401 2.51E−06 0.268464648 0.401149077 cg01533258 GSC N_Shelf Body 8.88002 1.32E−06 0.112399653 0.236846456 cg01550148 H2AFY S_Shore TSS1500 8.44036 0.000622 0.296437357 0.356835615 cg01564135 EVX1 N_Shelf TSS1500 6.00986 1.67E−05 0.446642618 0.594596172 cg01587682 PAX6 Island Body 11.8454 9.96E−12 0.252639806 0.474091567 cg01602621 ACP5 S_Shore TSS1500 6.85593 4.00E−06 0.355163159 0.481147742 cg01622304 PDX1 N_Shore Body 6.31336 0.000977 0.175900355 0.246920972 cg01627823 PAX9 N_Shore TSS1500 9.13267 6.10E−08 0.285641891 0.459152363 cg01631162 RNF39 Island 1stExon 8.45506 3.54E−06 0.209455381 0.33377347 cg01645998 SDR42E1 Island TSS200 14.7948 2.63E−08 0.396627711 0.570259373 cg01655831 GREB1L Island 5URT 8.18989 2.57E−06 0.174013931 0.291218337 cg01658421 EVX2 N_Shelf Body 7.32668 2.44E−05 0.318822401 0.428823948 cg01665555 PAX6 Island 5URT 3.52945 0.001039 0.324205235 0.424872836 cg01673674 GDF6 N_Shelf Body 11.0363 1.65E−06 0.457229408 0.597537303 cg01693350 WT1 Island 1stExon 8.16132 6.32E−07 0.425426445 0.59533214 cg01708273 HOXD11 S_Shore 3UTR 4.45151 0.00055 0.240404204 0.349318532 cg01713701 DLX4 S_Shelf TSS1500 4.05267 0.005361 0.312889498 0.382019266 cg01727145 SPEG Island Body 15.4120 5.56E−06 0.528836513 0.68062323 cg01739725 TTC6 N_Shelf 5URT 8.02241 3.23E−07 0.120323456 0.24486317 cg01789499 PRDM8 Island 5URT 8.77905 0.000104 0.598265837 0.688878343 cg01803928 DLEU1 Island Body 5.43855 0.000248 0.369220664 0.468627013 cg01811325 EBF3 Island Body 11.1627 7.47E−08 0.19962491 0.333947493 cg01815671 CCDX37 Island TSS200 8.24919 7.25E−08 0.101209468 0.236943776 cg01819512 HOXA10-AS N_Shore TSS1500 8.58434 0.000227 0.444329961 0.524834808 cg01821018 TACSTD2 Island TSS200 8.26762 0.000106 0.541982868 0.634825475 cg01824969 TCF24 Island TSS1500 6.80998 3.74E−05 0.153891159 0.24947656 cg01832036 DLX4 Island Body 6.4572 8.90E−07 0.214231841 0.385945834 cg01850449 LYPD1 Island 5URT 5.83717 0.004198 0.136623119 0.190500443 cg01853561 SIM2 Island 3UTR 11.1775 6.88E−07 0.381472052 0.525188129 cg01882880 HOXB-AS1 N_Shelf Body 6.34311 0.001074 0.645805907 0.722304869 cg01896761 FBLN1 Island TSS200 4.74647 2.44E−05 0.23701325 0.869284338 cg01904410 DKFZp686K1684 Island TSS1500 9.84664 2.90E−07 0.195901616 0.323569333 cg01913455 TBX2 Island Body 26.8418 3.05E−06 0.430191978 0.541895 cg01921432 BARHL2 Island 1stExon 16.8334 2.25E−10 0.123501363 0.246600247 cg01926269 POMC Island TSS200 7.22484 0.000243 0.289255916 0.386545049 cg01934626 SCG8SA1 Island TSS200 7.81221 1.42E−09 0.134863643 0.337329678 cg01939477 MIR129-2 Island TSS200 13.2881 4.20E−11 0.060656044 0.190832 cg01942863 GPC2 Island Body 13.0303 2.08E−09 0.370223887 0.554347334 cg01946451 NBPF8 S_Shore Body 11.1883 4.39E−08 0.250016988 0.374381152 cg01961447 GALR3 Island Body 11.9563 7.75E−09 0.07429998 0.209523388 cg01972418 PAX9 N_Shore TSS1500 7.62618 9.23E−07 0.2821947 0.418507289 cg02005600 HOXA5 Island TSS1500 4.79515 0.003763 0.479041426 0.548242707 cg02006615 PTPRN2 Island Body 5.48878 0.002594 0.498599567 0.569470677 cg02022733 PCDHGC4 Island TSS200 4.4107 6.00E−05 0.320379435 0.472483865 cg02032558 LRAT S_Shore 5URT 6.45159 0.000284 0.561049896 0.686101545 cg02037307 PITX1 Island 3UTR 9.79597 2.84E−08 0.383990581 0.581448187 cg02051616 NKX2-5 Island Body 14.8523 1.46E−07 0.328432884 0.444282737 cg02055963 CDX2 S_Shelf TSS200 10.1156 5.47E−08 0.214485474 0.345188768 cg02058624 LINC00403 N_Shore Body 6.21801 1.62E−05 0.254777874 0.380992929 cg02069715 GABRG3 Island TSS1500 7.85619 3.86E−05 0.346038304 0.467250822 cg02086801 OTP N_Shelf Body 7.20521 5.78E−05 0.413101419 0.504017667 cg02100373 PITX1 S_Shore Body 19.3543 6.29E−09 0.313121857 0.450246491 cg02102424 SYN2 Island Body 5.43408 0.00318 0.298312453 0.361996214 cg02108033 FAM1843 Island TSS200 11.2464 3.04E−07 0.072428905 0.165648982 cg02109484 M1AP Island TSS200 5.75708 2.76E−05 0.235539929 0.337535602 cg02115599 HOTTIP Island Body 4.94818 0.000277 0.136968084 0.224072282 cg02120658 PAX3 Island Body 5.59853 0.00093 0.149631291 0.216708531 cg02132714 HOX54 S_Shelf TSS1500 4.14282 0.000133 0.280682901 0.38808181 cg02145932 ISM1 Island TSS1500 9.45599 9.87E−08 0.10924131 0.229839915 cg02150354 NR2R1-AS1 N_Shore Body 10.0314 7.00E−08 0.338864078 0.548808297 cg02151625 PRR18 Island TSS200 6.90138 2.79E−05 0.199344395 0.30192599 cg02154252 ADGRL4 Island TSS200 9.17623 0.000115 0.31824814 0.414856202 cg02185185 RNF39 S_Shore Body 8.62369 0.001494 0.607472381 0.687571354 cg02194789 SIM2 S_Shore Body 4.6577 0.000271 0.16482024 0.262396915 cg02213684 PITX1 S_Shore Body 8.18273 1.42E−08 0.216359574 0.387903838 cg02233197 TNFAIP8L3 S_Shelf Body 4.60211 0.006792 0.564692736 0.633177768 cg02250594 ONECUT2 Island 1stExon 6.59273 2.28E−06 0.27143413 0.41810386 cg02281208 GABRG3 Island TSS1500 21.2525 1.28E−10 0.180299421 0.334801792 cg02283931 SDR42E1 N_Shore 5URT 7.32431 4.94E−05 0.512484868 0.627216747 cg02285920 JAK3 S_Shore TSS200 7.58997 0.000774 0.4946198 0.594739188 cg02285091 BHMT 5URT 93.552 4.65E−09 0.604337512 0.734554 cg02330683 ITPKA Island Body 13.2753 0.000165 0.62405414 0.736696455 cg02335804 HNF18 S_Shore TSS200 4.61338 0.001419 0.122732664 0.1975712921 cg02346492 NTN1 Island Body 3.64372 0.002117 0.184303237 0.264962073 cg02348751 LOC90768 Island Body 9.49702 3.01E−10 0.179731474 0.385594779 cg02360980 LTK Island Body 6.66606 1.57E−05 0.079910642 0.171702881 cg02364238 SHISA3 N_Shore TSS1500 9.71748 1.15E−06 0.158354457 0.256364945 cg02367696 LBX1-AS1 Island Body 10.49 1.72E−07 0.331959318 0.476934305 cg02428056 PAX9 S_Shore TSS1500 10.3867 3.07E−06 0.269526427 0.569440748 cg02435495 HNF1B Island Body 3.25385 0.001407 0.201856548 0.514692642 cg02439266 HOXA3 Island Body 6.91117 0.000324 0.575609287 0.660179293 cg02458062 HOXB3 S_Shore Body 5.2714 0.000541 0.412196209 0.493733 cg02468250 PAX6 Island 5URT 4.70147 0.000709 0.240163217 0.328508347 cg02485701 HOXA10-HOXA9 N_Shelf Body 7.07197 0.000511 0.491468411 0.558451404 cg02487130 SPEG Island Body 4.02835 0.002897 0.262993717 0.359651737 cg02495310 PITX1 S_Shore Body 10.8464 1.72E−09 0.230644126 0.416737188 cg02497558 HOXB1 S_Shore TSS1500 4.06605 0.000314 0.469568077 0.598702245 cg02511456 MNX1 Island Body 7.03104 2.97E−06 0.396138753 0.531196366 cg02524946 TBX20 Island TSS1500 14.2593 2.60E−08 0.185506974 0.301015227 cg02530022 DLX6-AS1 N_Shelf Body 5.85943 1.71E−05 0.26657213 0.395653072 cg02552311 RNF39 Island Body 6.81085 2.27E−06 0.082060426 0.196994138 cg02553516 SIM2 N_Shore Body 9.30088 3.16E−07 0.202993038 0.33832287 cg02569236 ALDH1L1 S_Shore 1stExon 6.96139 1.82E−05 0.158801595 0.254817317 cg02585702 BCAT1 N_Shore 1stExon 9.25237 5.22E−06 0.241930915 0.355081176 cg02595832 FOXI2 Island 1stExon 9.87276 1.44E−09 0.087475403 0.229388163 cg02596153 NR2E1 Island Body 4.05372 0.008121 0.203314488 0.263205132 cg02599464 HIST1H2BK N_Shore 3UTR 5.78193 4.37E−05 0.250990051 0.382206029 cg02611848 M1AP Island TSS1500 7.0688 6.38E−08 0.187664935 0.351880054 cg02623050 FAM84B Island Body 12.6305 3.48E−09 0.157291424 0.293563177 cg02624770 TRIL Island Body 10.4554 7.20E−08 0.326162386 0.4822965 cg02628323 MGAT4D S_Shore TSS200 5.79031 3.07E−05 0.277233507 0.385823818 cg02633371 LHX1 Island Body 7.3666 0.000109 0.334927605 0.416747636 cg02642123 PAX6 Island 5URT 6.76549 6.70E−07 0.204792677 0.347364245 cg02642822 HOXB-AS3 N_Shelf TSS1500 5.76502 0.000133 0.16358525 0.252092073 cg02645491 KCNQ1DN Island TSS1500 14.2004 1.60E−09 0.333097952 0.497769223 cg02655630 SLC12A5 Island Body 9.23931 7.90E−06 0.324876543 0.42836099 cg02660440 MIR199A1 TSS200 26.1824 1.72E−06 0.670980946 0.799887747 cg02688752 F2RL1 S_Shore Body 10.9516 7.79E−08 0.114254047 0.232900868 cg02694427 HOXD12 S_Shelf TSS200 10.9272 9.05E−11 0.240707797 0.427205006 cg02700894 SYN2 N_Shore TSS1500 5.45005 0.003265 0.183851555 0.301186263 cg02701080 SIM2 N_Shore Body 7.26188 8.85E−05 0.295690425 0.385924016 cg02703870 SDR42E1 Island TSS200 10.0695 1.10E−06 0.269207568 0.401618841 cg02741882 SEZ6L2 Island Body 23.1324 8.25E−07 0.378327236 0.513288477 cg02760031 TCF15 Island TSS1500 10.6235 9.06E−07 0.161988444 0.284281806 cg02767242 TBR1 S_Shore Body 4.97074 0.000486 0.495354667 0.815470581 cg02768694 M1AP Island TSS1500 8.02086 0.000225 0.455294558 0.543984071 cg02770983 PAX6 N_Shelf Body 17.2812 2.66E−08 0.491579357 0.646022615 cg02771142 PAX6 N_Shore Body 18.0344 5.98E−11 0.305157093 0.484222313 cg02781618 BEND4 Island Body 10.5191 4.97E−08 0.251525191 0.395463694 cg02798576 LINC01475 Island Body 5.77888 7.61E−05 0.111466728 0.197559236 cg02800607 HPSE2 N_Shore Body 4.90126 0.004655 0.149575929 0.220563177 cg02803914 APB81IP Island 5URT 15.6982 4.20E−09 0.1684828 0.289341751 cg02865068 DKFZp686K1684 S_Shore Body 6.01837 0.000616 0.514782295 0.607527608 cg02873421 HOXB3 S_Shore Body 4.84173 0.003725 0.425897457 0.495652283 cg02874942 ACP5 Island 5URT 8.81912 2.41E−08 0.13720768 0.273386218 cg02878244 DBX1 Island TSS200 8.67144 4.64E−07 0.123444715 0.231479873 cg02885007 HOXD9 Island 1stExon 13.7751 1.01E−09 0.134741609 0.264423439 cg02886033 HOXA10-AS N_Shore TSS1500 5.04793 0.006094 0.593816062 0.674513212 cg02906407 SSPO N_Shore ExonBod 47.4153 7.29E−08 0.701407643 0.856738889 cg02907064 MIR199A1 TSS200 11.5991 0.000343 0.656385047 0.755694485 cg02916932 PLLP Island TSS1500 5.35762 2.21E−05 0.150924617 0.282460365 cg02948476 GSC Island Body 6.05619 1.39E−05 0.160540538 0.27277727 cg02964031 COL14A1 Island 5URT 10.3867 1.92E−07 0.116243284 0.221621627 cg02971322 LBX1-AS1 S_Shore Body 74.3183 7.86E−09 0.569535519 0.690219899 cg03001942 LINC01143 S_Shore Body 5.66514 0.001736 0.173263916 0.229577945 cg03011535 TOX2 Island 1stExon 5.98231 0.000339 0.078471913 0.150466429 cg03020554 STK3 N_Shore 5URT 3.8879 0.00137 0.299884358 0.38961328 cg03024760 GABRG3 Island TSS1500 6.09499 5.71E−08 0.199487986 0.41034481 cg03056592 LINC01475 Island Body 8.28478 8.82E−07 0.13420514 0.23278097 cg03045635 DRD5 Island TSS200 10.2921 2.00E−09 0.179558772 0.350196017 cg03086235 SOX7 Island Body 9.36209 3.34E−05 0.473429494 0.594646455 cg03091551 OTX2 Island Body 12.5473 6.20E−08 0.176764776 0.279247115 cg03099728 ZNF274 S_Shore 5URT 17.3353 0.000116 0.728492667 0.814833091 cg03102494 NBPF8 Body 6.40069 1.40E−05 0.441430231 0.566673865 cg03129884 MAL S_Shore Body 6.67144 2.87E−06 0.191864533 0.298059662 cg03133868 NKX2-2 Island 3URT 6.64641 2.67E−05 0.359596481 0.476189303 cg03144922 PITX2 S_Shore Body 7.8105 3.87E−07 0.190462605 0.30788119 cg03148184 PITX2 S_Shore 5URT 4.10567 0.00189 0.241987271 0.327200451 cg03171465 KIAA1211L Island TSS1500 15.9753 4.75E−07 0.447094099 0.586707677 cg03177551 ITPKA Island Body 15.6101 2.77E−13 0.151671396 0.352050624 cg03181248 SP9 Island TSS200 26.0349 2.57E−09 0.150059892 0.26878577 cg03229282 RNF39 Island Body 7.77047 2.52E−06 0.08418392 0.190425975 cg03219968 ESRRG N_Shore 5URT 7.36827 8.61E−09 0.24347508 0.427494335 cg03242698 LBX1-AS1 S_Shore Body 19.043 4.08E−11 0.394950794 0.601149701 cg03276408 LINC00403 Island Body 8.99938 8.00E−08 0.094000168 0.222792543 cg03293507 RNF39 N_Shelf Body 10.5225 3.39E−05 0.59442824 0.700764162 cg03301582 PON3 Island TSS1500 13.0758 8.95E−09 0.278756802 0.411526847 cg03323462 LHX1 Island 3UTR 12.8563 4.64E−07 0.096686791 0.183477551 cg03325664 GATA4 N_Shelf TSS1500 5.27072 0.00869 0.274042393 0.363782997 cg03343571 RNF39 Island Body 5.27825 2.50E−05 0.319013513 0.456539884 cg03347590 PITX1 S_Shore Body 5.46476 1.87E−05 0.184870599 0.327529286 cg03350299 APOB Island TSS200 5.79666 0.000618 0.462482775 0.559319899 cg03355900 QRFPR Island TSS200 13.5323 1.26E−08 0.217274385 0.364720474 cg03405909 DKFZp686K1684 N_Shore Body 10.7846 4.08E−12 0.298723347 0.535053654 cg03405983 LYNX1 Island 5URT 5.73527 0.00163 0.416113124 0.500072263 cg03431846 REC8 Island TSS200 25.1931 4.38E−13 0.3025359 0.535462101 cg03465861 ISL2 Island Body 13.5627 1.92E−07 0.33617466 0.472074649 cg03491584 JAK3 S_Shore TSS200 8.38591 0.000139 0.350502716 0.453809687 cg03522132 LBX2 Island 3UTR 12.5129 1.85E−08 0.287045305 0.406859622 cg03529432 HOXA-AS3 Island Body 10.1972 2.77E−07 0.176740755 0.306479652 cg03562044 EPHX3 Island Body 14.6068 1.82E−12 0.098006067 0.267463674 cg03572772 PCDHB15 S_Shelf TSS200 4.87163 0.000123 0.118397759 0.206668348 cg03583111 DES Island 1stExon 20.5848 3.94E−11 0.197015481 0.332682502 cg03596280 GAD2 Island Body 4.20903 0.00444 0.12007581 0.178699508 cg05609639 LINC01475 S_Shore TSS1500 12.6355 6.78E−08 0.294703821 0.420858768 cg03636532 ONECOT2 Island Body 8.77141 2.55E−06 0.222343911 0.326343228 cg03637815 LOC101927248 Island Body 9.78757 3.72E−08 0.118046125 0.253463774 cg03640756 PCDHGC4 Island TSS200 6.27644 5.18E−06 0.231813034 0.362171047 cg03650674 GRHL2 Island Body 18.1371 6.00E−16 0.144644841 0.367358931 cg03655683 OTX1 S_Shore Body 17.4863 1.44E−11 0.261859764 0.446722446 cg03656099 TBX2 N_Shelf Body 3.73222 0.000653 0.180804303 0.289345535 cg03660500 LHX9 Island Body 5.775 0.000888 0.163508712 0.241639494 cg03669298 GATA3 S_Shore Body 8.03398 0.000257 0.406613802 0.520518353 cg03672342 GATA3-AS1 Island Body 3.32123 0.002435 0.212879417 0.317958259 cg03692563 DKFZp686K1684 N_Shore Body 13.9572 8.98E−11 0.208013049 0.365422262 cg03694718 NKX2-6 Island TSS200 17.021 4.16E−08 0.129976099 0.232864124 cg03695236 PTPRN2 Island Body 7.21056 0.00012 0.470650501 0.55757427 cg03711485 NKX2-3 N_Shelf 3UTR 6.60456 9.83E−07 0.341733319 0.485493122 cg03733219 SPRED3 Island Body 5.17645 0.001083 0.300898205 0.373052289 cg03733760 FGF19 S_Shore Body 4.54743 0.000172 0.256462395 0.369348746 cg03735496 GREB1L Island 5URT 8.25656 2.05E−07 0.079158539 0.197425684 cg03744440 MYO15B Island TSS200 18.6841 4.95E−19 0.140396281 0.337035329 cg03744763 HOXA5 Island TSS1500 10.5054 1.19E−06 0.237168054 0.327892148 cg03771840 TRIM15 Island 3UTR 11.2579 0.002406 0.711416457 0.780360899 cg03774463 DLX6 S_Shore Body 5.2222 0.000234 0.28273975 0.37267829 cg03780851 GRHL2 Island TSS200 10.2671 6.65E−08 0.270769599 0.44580223 cg03826976 CYB5R2 Island 5URT 10.8637 1.95E−05 0.452616725 0.567828804 cg03851159 VAX1 N_Shore 3UTR 7.04373 4.90E−05 0.142264788 0.241347655 cg03854796 PAX6 N_Shore Body 16.0162 1.46E−07 0.395560566 0.517687525 cg03867465 LOC90768 Island Body 9.10674 2.48E−06 0.479714023 0.651848155 cg03885037 ADAMS Island Body 5.28413 0.000205 0.236108988 0.343000682 cg03905867 PAX6 Island Body 11.8021 1.15E−10 0.264501852 0.469462673 cg03927133 ITPKA Island Body 23.5197 9.65E−11 0.262750464 0.416308498 cg03945700 PRR18 Island TSS200 6.96745 0.00022 0.217055362 0.305625365 cg03947464 UPK3A Island TSS200 10.5551 5.67E−08 0.112067382 0.228035911 cg03951853 NBPF8 Body 7.06289 0.000235 0.445971867 0.501907495 cg03958979 NR2E1 Island TSS1500 9.33962 2.28E−05 0.376855736 0.475414889 cg03959515 FAM84B Island 5URT 6.47902 2.88E−07 0.17301173 0.523691305 cg03960072 KAAG1 Island 1stExon 36.9545 8.11E−10 0.314402351 0.439506303 cg03964958 HOXD12 Island 1stExon 12.5171 5.30E−09 0.122362503 0.245863702 cg03970615 IRX3 Island Body 7.28154 3.63E−05 0.308284913 0.426766852 cg03975235 GALR3 Island Body 11.5382 0.000824 0.489935519 0.567151626 cg03981954 FEZF1 N_Shore 3UTR 5.81951 1.31E−06 0.149783744 0.274631337 cg03989480 DMRTA2 Island Body 5.66812 0.000182 0.385058427 0.428202706 cg04022561 LOC642366 Body 11.2364 4.34E−08 0.114521412 0.231744442 cg04025675 GATM Island TSS200 3.55026 0.004475 0.220640217 0.307631915 cg04046669 HORMAD2 Island TSS1500 13.1346 1.15E−12 0.179208873 0.403051353 cg04047221 SOX1 Island 3UTR 12.2824 1.71E−08 0.146817957 0.273742583 cg04052466 AMH Island Body 8.48992 2.13E−09 0.246444269 0.446683465 cg04059568 PTPRN2 Island Body 8.56297 6.16E−07 0.154539162 0.268259715 cg04065086 SEMA6C Island 3UTR 5.05 3.87E−05 0.383561956 0.528837553 cg04079760 ESRP2 N_Shelf Body 23.3056 1.03E−09 0.451476727 0.628448162 cg04080041 PCDHGA12 Island 1stExon 12.8918 8.72E−09 0.331951211 0.500179594 cg04080282 PON3 Island TSS1500 24.2223 7.66E−08 0.527982992 0.66176999 cg04091063 AKR7A3 Island TSS200 15.0349 1.51E−08 0.296122638 0.475935021 cg04131898 WT1-AS Island Body 5.56518 0.001755 0.1314334 0.188783143 cg04134048 TNFRSF10D S_Shore TSS200 7.28106 7.85E−06 0.347644368 0.493013443 cg04136369 HNF1B Body 27.3908 1.82E−05 0.687851837 0.796073717 cg04142323 LINC00461 S_Shore TSS1500 9.88372 5.47E−06 0.18544802 0.285239913 cg04158367 GFI1 Island TSS1500 8.70001 1.01E−06 0.301580489 0.452683437 cg04178316 TLX3 N_Shelf Body 5.23369 0.001939 0.157414454 0.228749578 cg04180086 IRX4 Island Body 8.87726 4.74E−06 0.145040065 0.239534967 cg04193970 TTC6 Island Body 9.63885 2.56E−10 0.181277708 0.377229253 cg04198914 HNF1B S_Shore TSS1500 3.37251 0.005973 0.427471109 0.501742755 cg04203645 APC2 Island Body 5.44756 0.000626 0.407663692 0.508755421 cg04213746 GATA3 Body 7.14256 0.001142 0.721690977 0.808597889 cg04236178 ESRRG N_Shore 5URT 10.3024 1.19E−08 0.329913803 0.490934079 cg04251368 KCNQ1DN Island TSS1500 7.1927 2.99E−07 0.195311103 0.343130074 cg04255230 LBX2 Island Body 4.81047 4.45E−05 0.271341088 0.415139664 cg04265576 HOXA-AS3 Island Body 7.48154 6.92E−05 0.133919803 0.206062225 cg04290346 D6X1 Island Body 8.87731 1.90E−07 0.096948607 0.202125059 cg04328477 ESRP2 Island TSS200 4.10116 0.002371 0.372415165 0.462467045 cg04342594 LINC01391 N_Shore TSS200 9.22501 1.00E−05 0.415627669 0.527453626 cg04360793 ADGRL4 Island 1stExon 5.17727 1.14E−06 0.34204361 0.537357309 cg04366687 OPLAH Island Body 14.2546 1.96E−06 0.349784004 0.467116788 cg04369341 TOX2 Island Body 4.74436 0.003488 0.215849031 0.288371991 cg07378886 RCN3 S_Shore 5URT 10.0352 1.56E−07 0.27244142 0.423281952 cg04403809 SLC12A5 Island 3UTR 27.1545 1.05E−07 0.28116514 0.374360717 cg04407470 NR2E1 Island Body 5.06396 0.000235 0.060659067 0.185995183 cg04415176 HOXD13 Island 1stExon 9.12268 3.49E−06 0.083375797 0.191715974 cg04415798 PAX9 S_Shore 5URT 4.27388 4.18E−05 0.152159215 0.287995812 cg04418091 ATP5G2 Island TSS1500 3.59509 0.001164 0.14083792 0.24525696 cg04446345 UPK3A Island TSS200 8.81311 1.58E−05 0.208272716 0.312046441 cg04450456 FAM1843 Body 17.4558 6.86E−08 0.557261791 0.687451505 cg04456238 WT1 S_Shore Body 11.4207 3.82E−07 0.336912504 0.465736645 cg04461228 NR2F1-AS1 Island Body 13.8976 2.55E−07 0.424953536 0.56465222 cg04467589 RNF35 S_Shore TSS1500 11.9626 0.000351 0.445556775 0.505938091 cg04469219 LINC00403 Island TSS1500 11.6448 2.27E−08 0.091970592 0.150622294 cg04489012 C2CD4D S_Shore 5URT 12.3436 4.19E−06 0.456766573 0.598791375 cg04504066 PAX6 S_Shore 5URT 20.5935 1.58E−11 0.180852284 0.338366544 cg04513006 ESRP2 Island TSS200 3.7056 0.004063 0.286814628 0.357666594 cg04514123 SYNGR8 S_Shore Body 10.8376 7.89E−10 0.145986502 0.300849631 cg04515001 KAAG1 Island 1stExon 6.55596 3.75E−08 0.186840236 0.354998889 cg04521510 HOTTIP N_Shelf Body 5.00434 0.000169 0.125783385 0.221535142 cg04543413 BCAT1 Island TSS1500 16.1327 1.21E−09 0.184927566 0.317019285 cg04552206 TLX1 Island Body 6.62135 0.000726 0.273026248 0.346458455 cg04554928 TRIM15 Island 3UTR 6.35467 0.002373 0.478086349 0.564725768 cg04558175 RCN5 S_Shore 5URT 11.7018 6.74E−05 0.628132868 0.748755818 cg04566159 ACP5 Island 5URT 11.3638 4.50E−05 0.402818565 0.536090131 cg04574054 PRAC2 S_Shelf TSS1500 4.51916 7.96E−05 0.291220253 0.419839379 cg04586579 SDR42E1 Island 5URT 8.09315 5.29E−06 0.258990266 0.397663561 cg04597433 DRD5 Island TSS200 7.81426 1.40E−05 0.119448781 0.216242634 cg04597449 GPR150 N_Shore TSS1500 3.43764 0.008851 0.333395374 0.413057141 cg04598774 PAX6 S_Shore Body 27.3606 8.60E−13 0.218210078 0.38766699 cg04617948 FOX12 Island TSS1500 31.6937 7.76E−10 0.43486481 0.6017015 cg04623837 HCG9 N_Shore Body 7.98941 4.53E−07 0.298498815 0.480692037 cg04630292 PRRT1 Island 3UTR 3.98086 0.006055 0.459294058 0.504342645 cg04636269 LHX1 Island 3UTR 5.20672 0.001157 0.812285322 0.39083719 cg04648480 SIM2 S_Shore Body 5.27688 0.000265 0.401238405 0.502791257 cg04654580 OTX1 Island Body 9.48413 1.97E−10 0.275466332 0.475796179 cg04673325 ZNF518B Island TSS200 10.5496 1.76E−08 0.152019918 0.281781773 cg04685170 PON8 Island TSS200 5.98358 0.00024 0.309790053 0.387507674 cg04688051 LYNX1 Island 5URT 6.72173 2.46E−07 0.142443015 0.285497693 cg04728480 APBB1IP N_Shore TSS1500 3.59417 0.001725 0.256159391 0.337756086 cg04729913 GATA3-AS1 Island Body 12.3779 0.00013 0.465038411 0.556351222 cg04730685 LOC101927248 S_Shelf Body 14.9126 1.28E−07 0.381836222 0.514076535 cg04738888 MYO15B Island TSS200 30.5145 1.18E−12 0.25561768 0.427184443 cg04739647 HCXD9 Island 5URT 6.64276 2.96E−05 0.149606771 0.248272855 cg04765277 GATA3-AS1 Island Body 4.89144 0.000145 0.292924556 0.419140007 cg04775710 IERS Island Body 12.7649 3.30E−09 0.214689078 0.389077326 cg04787024 KIAA1211L Island TSS1500 8.21827 5.20E−10 0.268113979 0.496846403 cg04796181 HIST1HB1 Island 1stExon 11.4896 1.19E−08 0.133404567 0.264818755 cg04809787 CHRNB1 Island TSS200 10.9969 9.78E−07 0.249933354 0.371353219 cg04821708 SDR42E1 Island TSS200 12.5748 8.20E−07 0.333422556 0.467231894 cg04850999 C10orf11 Island Body 7.87629 0.003061 0.713162171 0.801042697 cg04854328 NR2F1-AS1 Island Body 6.47623 1.80E−05 0.140541447 0.259469238 cg04858858 AGAP1 Island TSS1500 8.55341 4.81E−08 0.25187456 0.42131318 cg04864199 SHF S_Shelf 5URT 30.8248 5.11E−09 0.497527629 0.678207076 cg04886060 SDR42E1 N_Shore 5URT 8.56277 8.41E−06 0.473221262 0.615325889 cg04887494 JSRP1 Island Body 4.30211 3.51E−05 0.521180662 0.470707462 cg04904318 HOXB1 Island 1stExon 4.05689 0.001389 0.517519174 0.612871879 cg04916091 JAK3 Island 5URT 5.67689 4.51E−05 0.256277892 0.396706996 cg04937416 PTPRN2 Island Body 3.3275 0.004443 0.442201133 0.532210548 cg04938549 PAX6 N_Shore 5URT 10.3475 4.49E−13 0.220136969 0.488327634 cg04956913 IER3 Island TSS200 7.3777 2.80E−05 0.256258891 0.398895634 cg04959835 CYB5R2 Island 5URT 7.36015 8.67E−06 0.11218199 0.210789062 cg04972566 FGF19 N_Shore Body 5.95737 7.26E−06 0.185602878 0.302986239 cg04972745 LINC01475 Island Body 4.01885 0.000676 0.813178154 0.415709533 cg04996219 CTNND2 Island 1stExon 9.46443 4.89E−08 0.068156759 0.17834887 cg05001964 IRX3 Island Body 4.25429 0.000137 0.158731104 0.285621993 cg05009601 FEZF1 Island 1stExon 7.70169 4.62E−06 0.309059788 0.436804884 cg05027336 HOXA10-A5 Island Body 5.82266 0.008025 0.142675695 0.208350182 cg05029822 KCNQ1DN Island TSS200 14.6722 3.63E−05 0.318111847 0.390930719 cg05031521 LMF1 Island Body 11.4525 9.96E−07 0.183132995 0.312258364 cg05065507 TACSTD2 Island 1stExon 17.6008 1.21E−07 0.235009493 0.325875895 cg05072100 LYNX1 Island 5URT 7.90314 1.16E−06 0.209027203 0.335639659 cg05140806 SHH N_Shelf Body 5.19567 0.000604 0.177646875 0.266475565 cg05142617 MAL N_Shore TSS1500 9.20848 9.90E−07 0.393394839 0.541751609 cg05142765 GSC Island 3UTR 4.85842 0.000325 0.288989178 0.395936251 cg05158615 NPY Island TSS1500 7.534 2.45E−07 0.293681987 0.447260552 cg05165940 SATB2 Island 1stExon 11.2209 1.21E−06 0.28832389 0.382720922 cg05167251 HOXD9 Island 5URT 7.55624 2.51E−06 0.198387284 0.509700661 cg05180443 CHAD Island TSS1500 3.91693 0.000389 0.22316233 0.34412746 cg05193369 CYP26C1 N_Shelf Body 5.91729 0.000789 0.187634088 0.258500834 cg05196969 ISL2 N_Shore TSS1500 9.98068 4.12E−09 0.154595415 0.512671802 cg05210501 HOXA10-HOXA9 Island Body 6.91782 0.000241 0.195591357 0.263659144 cg05222924 WT1 N_Shore Body 6.44924 1.41E−05 0.088734745 0.185265833 cg05236677 STK3 Island 5URT 8.32458 2.80E−06 0.224798536 0.358093791 cg05256269 HOXO13 S_Shore TSS1500 16.8971 3.03E−08 0.304113184 0.435109899 cg05290058 KCNQ1DN Island TSS1500 5.06864 1.37E−05 0.410864329 0.561331284 cg05301866 TBR1 Island Body 22.5958 2.23E−08 0.410505628 0.518827919 cg05317714 KIFC2 Island Body 57.2249 4.78E−14 0.295231785 0.529090365 cg05345154 AMH Island Body 0.01784 1.86E−09 0.24790974 0.459142351 cg05347898 KAAG1 Island 1stExon 6.90488 6.57E−08 0.096868476 0.2423145 cg05351827 SPEG Island Body 14.5346 6.51E−05 0.533432313 0.643321309 cg05368740 HSPB6 Island Body 4.28673 0.001806 0.328825281 0.420606382 cg05376374 MIR129-2 Island TSS200 15.6956 1.05E−12 0.059778181 0.203785349 cg05377576 SEZ6L2 N_Shore Body 16.7414 6.58E−05 0.554689665 0.670872374 cg05380821 NR2E1 Island Body 5.1387 0.000716 0.236612465 0.313793468 cg05387167 HOXB3 S_Shore Body 8.21026 0.000217 0.456289597 0.529069051 cg05406649 HOXO4 Island 5URT 4.35073 0.006925 0.738861209 0.785422275 cg05449136 PRO31 Island 1stExon 11.3355 5.93E−10 0.146969409 0.313063182 cg05457563 APC2 Island Body 6.97355 0.000893 0.376479927 0.462124805 cg05470554 WNT16 N_Shore Body 8.54167 2.53E−09 0.235299112 0.437242842 cg05475934 AP6B1IP Island 1stExon 7.39348 4.52E−07 0.186522658 0.329101564 cg05477444 INPP5A Island Body 11.215 0.000671 0.714821558 0.789695202 cg05488632 EPHX3 Island 1stExon 85.8683 1.03E−12 0.390514822 0.559915543 cg05490659 HOXA10 S_Shelf Body 4.21112 0.001722 0.245250948 0.335412057 cg05498007 KATNAL2 N_Shore TSS200 6.58294 8.12E−08 0.203589678 0.386896059 cg05500840 HOXD11 Island TSS200 12.527 1.38E−09 0.257812474 0.440698546 cg05501996 LOC642865 Body 14.5033 6.77E−13 0.184712326 0.374705466 cg05520409 LMF1 Island Body 21.1912 4.54E−06 0.358776165 0.305580327 cg05522011 PRDM8 N_Shore Body 3.07765 0.006963 0.624171176 0.708011343 cg05527530 LHX9 S_Shore Body 5.19586 0.002405 0.168759788 0.240135155 cg05529506 KATNAL2 Island 1stExon 8.19081 1.18E−08 0.137086846 0.311434044 cg05555207 TBX5-AS1 Island Body 5.48129 5.24E−05 0.148260033 0.232468557 cg05578840 OTP N_Shore Body 6.42376 7.65E−06 0.353794321 0.498461196 cg05592035 CD300A Body 9.31261 1.64E−07 0.308964828 0.486762482 cg05603881 SHISA3 S_Shore Body 4.78043 0.000903 0.147096058 0.217968074 cg05604079 QRFPR Island TSS200 7.33625 2.96E−08 0.19745714 0.377169215 cg05635754 JAK3 Island 5URT 5.76455 8.06E−05 0.372507206 0.501759228 cg05639205 SHH Island Body 11.1548 1.36E−07 0.213370517 0.338072454 cg05656180 KCNQ1DN N_Shore TSS1500 4.58423 0.00978 0.329915253 0.397065605 cg05658487 IRX4 Island TSS1500 20.1532 1.34E−09 0.223245993 0.348368599 cg05667348 VAX1 Island Body 5.18024 1.19E−05 0.180764685 0.304268554 cg05678749 GABRG3 Island Body 5.68088 0.000174 0.136910791 0.228655153 cg05690644 GDF6 Island Body 53.7262 1.18E−07 0.597656312 0.772548253 cg05697849 ELAVL4 Island 5URT 19.5073 1.52E−09 0.275810197 0.404820727 cg05724997 BCL2 S_Shelf TSS1500 6.69085 1.37E−05 0.252322091 0.373295122 cg05736768 DFYS Island 1stExon 23.949 1.54E−12 0.187180757 0.340324042 cg05744332 MNX1 N_Shore Body 22.6507 5.49E−05 0.602450375 0.69059104 cg05766510 PTPRN2 Island Body 3.08586 0.007926 0.580438012 0.645238596 cg05787556 TLX3 Island TSS1500 20.4256 2.00E−10 0.217448751 0.345871152 cg05818685 CTNND2 Island Body 10.1471 0.000186 0.459162729 0.55107703 cg05822926 ICA1 Island TSS1500 23.2354 1.64E−08 0.338808268 0.479127516 cg05835105 PITX2 S_Shelf Body 9.07141 7.36E−12 0.203399107 0.452202498 cg05839235 NPR3 Island 1stExon 15.1652 9.97E−13 0.143704891 0.326434855 cg05853632 RNF39 Island 1stExon 6.44474 5.25E−06 0.218287822 0.352576493 cg05890484 BHMT TSS200 20.8374 1.03E−05 0.540446093 0.643199364 cg05896902 GATM Island TSS200 3.66491 0.004096 0.233615904 0.330899056 cg05899618 GDF7 Island TSS1500 13.4991 9.88E−09 0.205282742 0.35049826 cg05905176 TNFAIP8L3 Island Body 5.19304 2.05E−07 0.239630407 0.440286099 cg05915293 SYNGR3 S_Shore Body 5.921 2.27E−06 0.268004435 0.446662949 cg05921905 HOXA2 N_Shelf TSS200 4.78279 0.000329 0.134393118 0.224593638 cg05922610 NR2E1 N_Shore Body 19.7272 1.15E−09 0.230347274 0.341964081 cg05924583 TP73 Island 5URT 11.7884 4.17E−05 0.440948772 0.564297823 cg05925327 LOC101929076 Island Body 4.32618 0.000161 0.256723886 0.380661308 cg05928186 HOXA-AS3 Island Body 8.71673 0.000281 0.244843558 0.808103502 cg05929882 TBX5-AS1 Island Body 8.41999 4.00E−06 0.097954926 0.181912476 cg05937630 AKR7A3 Island TSS1500 7.98694 1.72E−07 0.178576405 0.353656995 cg05942128 HOXD11 Island TSS200 20.617 7.31E−11 0.29540255 0.448715437 cg05974274 PRDM8 N_Shore Body 8.18823 3.52E−07 0.197495261 0.390610299 cg05975410 LOC648987 S_Shore TSS1500 5.08264 0.001893 0.292908925 0.390920108 cg05976500 NTN1 Island Body 27.781 1.80E−15 0.164488905 0.696399184 cg05977669 HOXA11-AS Island Body 26.6529 1.39E−06 0.474222062 0.574849141 cg05990544 HIST1HB1 S_Shore TSS1500 10.5824 9.88E−11 0.135833619 0.297756112 cg06005695 HIST1HB1 S_Shore TSS1500 22.9412 9.73E−10 0.223187505 0.349656841 cg06023345 FAMB4B N_Shore 3UTR 7.26993 5.62E−05 0.359474059 0.492969881 cg06049868 LINC00403 S_Shore Body 7.8033 0.000157 0.291253663 0.378116404 cg06055873 HOXA2 N_Shelf 1stExon 16.631 1.01E−09 0.368348872 0.51710603 cg06060135 IRX1 Island Body 6.39132 1.40E−06 0.106804028 0.224072525 cg06102419 LOC151174 N_Shore Body 33.332 7.51E−06 0.579995612 0.675802646 cg06133205 DLEU1 N_Shore Body 5.58808 4.03E−05 0.188124649 0.292899905 cg06142537 LINC01143 Island TSS1500 5.4592 0.000749 0.224659264 0.304354166 cg06145234 SHH S_Shore 5URT 5.08883 3.27E−05 0.296388516 0.487446254 cg06148812 TBX1 Island 5URT 23.1023 1.18E−05 0.579195643 0.692901162 cg06150468 BATF3 S_Shore TSS1500 8.86271 7.67E−08 0.106960341 0.253960487 cg06157318 APBB1IP Island TSS200 10.1179 1.04E−06 0.186548505 0.313339229 cg06159394 DMRTA2 Island 5URT 5.49198 0.006272 0.526456171 0.576542889 cg06166809 GPR150 Island 1stExon 23.9557 5.28E−10 0.152129994 0.279860984 cg06178563 NKX2-2 Island TSS200 11.9168 1.25E−06 0.196263349 0.300952132 cg06198998 KATNAL2 N_Shore TSS200 6.99904 8.61E−09 0.141469797 0.318931845 cg06226158 LBX2 Island 1stExon 5.16483 0.000356 0.190362384 0.286278345 cg06228630 ZAR1 Island Body 25.2837 1.57E−12 0.383122488 0.555957939 cg06230843 FAMB4B Island Body 10.5875 1.52E−07 0.240957951 0.385725809 cg06238409 MECOM Island TSS200 4.04426 0.004556 0.161929971 0.228024847 cg06289355 NPR3 Island Body 9.85931 8.67E−09 0.183383193 0.346782453 cg06249604 RNF39 Island Body 3.93003 4.25E−05 0.263301856 0.415649398 cg06254440 LHX2 Island Body 3.95656 0.003764 0.211087295 0.287557315 cg06255601 DLEU1 Body 6.07287 8.25E−06 0.339033453 0.527471672 cg06262436 ISM1 Island TSS1500 10.3694 2.52E−07 0.223783874 0.34915112 cg06272038 ASCL2 Island TSS1500 12.661 1.07E−07 0.05079888 0.148702194 cg06274159 ZFP42 Island TSS200 9.81907 3.55E−11 0.201375337 0.41292663 cg06287951 DLX4 S_Shore Body 14.6705 2.10E−06 0.383564258 0.499547 cg06303875 CDX2 S_Shelf TSS200 11.0416 7.52E−08 0.182710151 0.295716758 cg06312283 PAX6 S_Shore TSS1500 19.2436 1.77E−12 0.137502719 0.270944685 cg06323023 HIST1HB1 S_Shore TSS1500 10.6927 4.52E−08 0.167164061 0.311430898 cg06367117 ALDOC 5URT 36.9295 3.47E−09 0.156168537 0.26086272 cg06367511 PLLP Island Body 6.80999 3.61E−06 0.40393705 0.544994926 cg06375967 ISL2 Island Body 6.26597 1.28E−05 0.19262367 0.282325826 cg06376715 TP79 Island Body 20.1013 5.64E−06 0.601157636 0.716308788 cg06382344 TBR1 Island Body 18.5958 3.72E−15 0.118598433 0.356284372 cg06382559 TLX1 Island Body 3.45764 0.008023 0.261063491 0.314466023 cg06386307 SIM1 Island Body 7.05704 4.02E−05 0.380077302 0.492924949 cg06390536 MYO158 Island TSS200 10.5771 2.08E−07 0.171430236 0.259146776 cg06410191 LVRN Island 1stExon 15.9907 7.04E−08 0.227524578 0.34438569 cg06410537 TLX3 Island 1stExon 16.4501 4.38E−09 0.155685643 0.29692154 cg06425620 PCDHGC4 N_Shore TSS1500 4.9205 5.88E−05 0.234660733 0.380898562 cg06440543 PRDM8 S_Shore 5URT 11.1971 2.68E−05 0.632337047 0.71132196 cg06451900 LOC145845 N_Shore Body 5.72026 0.00032 0.412754411 0.513207667 cg06488443 TBR1 Island Body 30.0459 2.94E−11 0.253759163 0.384046656 cg06490225 LHX1 Island Body 5.79275 0.00032 0.25866663 0.543269871 cg06497848 VAX1 Island Body 8.04193 0.000134 0.389835358 0.462445566 cg06520675 LBX1-AS1 Island Body 7.94911 1.85E−05 0.166851958 0.330531853 cg06585993 APBB1IP Island TSS200 7.00562 1.95E−05 0.146532671 0.245046367 cg06554090 LOC101929076 Island Body 8.21623 1.39E−06 0.28802705 0.423138456 cg06571387 HOXD12 S_Shore TSS1500 13.6857 1.20E−06 0.37625662 0.488498677 cg06579271 PLLP Island TSS1500 9.58144 3.05E−05 0.217289791 0.321234876 cg06615152 AMH Island Body 11.8308 2.85E−06 0.510913961 0.631844475 cg06626599 KCNQ4 Island Body 14.1385 1.53E−05 0.478628428 0.582496929 cg06632207 ATP5G2 S_Shore TSS1500 3.38557 0.008113 0.082155288 0.147204271 cg06637517 DNAH1 Body 4.35003 2.91E−05 0.291351881 0.458736311 cg06644373 SATB2 Island 5URT 6.49707 0.007679 0.28295666 0.364917125 cg06666008 PAX6 N_Shore Body 12.1661 9.51E−13 0.205925812 0.446187033 cg06667574 IGLON5 Island Body 8.20214 0.000784 0.63913838 0.745123697 cg06702880 LRAT N_Shore 5URT 6.28077 6.33E−06 0.118959668 0.22326278 cg06704518 GATA4 Island 5URT 4.59876 0.002197 0.207443826 0.28668601 cg06705930 PAX6 N_Shelf 5URT 9.69726 0.000335 0.630173628 0.727324172 cg06708634 SIM1 S_Shore Body 7.53796 1.93E−06 0.398001261 0.546436879 cg06710082 HCG9 N_Shore Body 6.90499 8.02E−07 0.371448774 0.560210055 cg06723856 ESRF2 Island 1stExon 6.99852 5.27E−05 0.065670505 0.134618864 cg06724305 DMRTA2 Island 5URT 4.293 0.005039 0.29089591 0.353960642 cg06732395 SCG33A1 Island TSS1500 22.6843 1.99E−10 0.27929364 0.426210253 cg06754197 MIR199A1 TSS200 35.8724 3.79E−06 0.614658202 0.719448232 cg06765217 TTC6 Island Body 10.2106 3.67E−11 0.071472511 0.215188256 cg06779449 TOX2 Island TSS200 10.7538 6.53E−07 0.080850172 0.185610946 cg06804210 CD8A S_Shelf TSS1500 14.3491 2.10E−07 0.33357462 0.45325798 cg06813578 TNFAIP8L3 Island Body 5.89644 5.70E−06 0.173513354 0.31028461 cg06818052 NTN1 Island Body 11.8809 4.34E−05 0.461386946 0.567006865 cg06818777 CHAD Island TSS1500 4.58976 0.000862 0.272921576 0.384367439 cg06825142 DRD4 Island TSS200 7.48677 5.40E−06 0.10020074 0.20381938 cg06842954 PITX2 N_Shore 5URT 7.23515 4.42E−07 0.180063419 0.304475343 cg06845853 WT1-AS S_Shore Body 10.6891 7.20E−12 0.149371217 0.322156318 cg06584084 NKX2-3 N_Shore TSS200 8.04585 0.000122 0.148501078 0.191719088 cg06866686 GSC S_Shore TSS1500 6.16697 0.001813 0.421445349 0.496426152 cg06872257 FGF20 S_Shore 1stExon 25.2623 3.76E−08 0.467465767 0.609856904 cg06911121 TBX5-AS1 Island Body 5.30745 0.000656 0.158440819 0.229145194 cg06916239 PAX3 S_Shelf Body 6.62476 7.97E−06 0.243614275 0.364036427 cg06942183 HOXB-AS1 S_Shore Body 4.34517 0.003037 0.284047372 0.364007342 cg06942701 TBR1 Island Body 86.3924 9.76E−13 0.407200876 0.543208242 cg06943420 MMP23B Island TSS1500 8.07491 3.97E−10 0.2100618 0.424244613 cg06962944 TOX2 Island TSS200 7.04067 2.54E−05 0.082193076 0.212878265 cg06966811 OTX2 N_Shore Body 16.9613 2.12E−10 0.31196753 0.492314091 cg06975043 KCNAB3 Island 3UTR 9.91146 4.79E−06 0.199350688 0.304158052 cg06991300 QRFPR Island 1stExon 13.819 7.38E−08 0.140336902 0.259937392 cg06992285 STK3 Island 5URT 5.37559 8.80E−06 0.216394633 0.35785876 cg07010088 SKOR1 Island Body 4.93113 0.003819 0.238633922 0.293784595 cg07010228 JSRP1 Island Body 8.90391 0.000163 0.255354814 0.332621396 cg07017875 PCDHG86 Island 1stExon 16.004 6.93E−07 0.479779496 0.598656949 cg07020001 KDM2B S_Shore TSS1500 4.86584 0.000879 0.143982427 0.239040424 cg07021644 UPK3A Island 5URT 10.2246 4.18E−07 0.226893244 0.351062474 cg07029873 ESRRG Island 1stExon 9.56191 3.69E−10 0.138112815 0.311573376 cg07033624 LINC00403 N_Shelf Body 14.0182 2.87E−08 0.164622669 0.282675488 cg07046818 GDF6 N_Shelf Body 6.04561 2.33E−05 0.348011051 0.463268873 cg07050616 LBX1-AS1 Island Body 9.62835 1.58E−08 0.157130462 0.318356733 cg07060233 SLC12A5 Island 3UTR 44.5703 1.88E−11 0.366929124 0.505961182 cg07095230 TBX2 Island Body 4.03414 0.000182 0.383930524 0.512640611 cg07104660 TCF24 Island Body 17.8474 2.87E−11 0.16164926 0.304270466 cg07116997 HOXA11-AS S_Shore Body 8.44641 0.000105 0.183230529 0.255319415 cg07121856 PON8 Island TSS200 15.2066 5.30E−13 0.141504037 0.334556926 cg07124117 PAX6 Island Body 12.8326 1.55E−09 0.219689223 0.382253517 cg07135614 LINC02391 N_Shore TSS1500 5.5274 9.57E−05 0.29196725 0.407797311 cg07147475 MGAT4D S_Shore TSS200 10.342 8.47E−06 0.395251711 0.518101239 cg07155223 ESRRG N_Shelf 5URT 5.78903 1.01E−05 0.09639401 0.199869088 cg07160932 SEZ6L2 S_Shore Body 9.21759 2.14E−08 0.286851089 0.502690903 cg07178825 TP73 Island 3UTR 49.6557 1.47E−13 0.440568326 0.619546192 cg07195011 CTNND2 Island 1stExon 8.80107 1.39E−09 0.085902935 0.229889145 cg07201017 LBX1-AS1 Island Body 12.5843 6.43E−08 0.325290793 0.46231202 cg07211140 WT1 S_Shelf Body 8.22282 2.03E−07 0.168148605 0.307013632 cg07247419 NkX2-4 Island 3UTR 12.7573 8.58E−07 0.315247287 0.434596404 cg07274716 PITX1 Island 3UTR 10.7107 1.03E−11 0.248936197 0.470384833 cg07281879 WT1-AS Island Body 11.8386 7.57E−08 0.137990289 0.255356689 cg07300846 SEZ6L2 S_Shore Exon3nd 5.76948 1.47E−05 0.254227504 0.408846359 cg07302910 VAX1 N_Shore 3UTR 4.40459 0.000651 0.265083289 0.35850637 cg07311313 COX2 S_Shelf TSS200 8.12688 1.09E−08 0.094611722 0.226825864 cg07322064 PRR18 Island 3UTR 11.0121 6.08E−05 0.466917442 0.549986838 cg07323141 LOC151174 Island Body 4.91533 0.002033 0.208585112 0.295466371 cg07329360 DKFZp686K1684 Island Body 16.6189 3.27E−11 0.312894591 0.480529035 cg07330230 ALDH1L1 S_Shore 1stExon 4.94285 3.63E−05 0.153605512 0.272795436 cg07333715 ASCL2 Island TSS1500 9.24636 3.62E−10 0.163843346 0.361614896 cg07359638 LTK Island Body 6.68189 0.000291 0.290410173 0.36553508 cg07367232 KCNA33 Island 3UTR 7.44099 1.03E−05 0.105800144 0.202783451 cg07371589 SEMA6C Island 3UTR 3.8852 0.000311 0.543665056 0.661379807 cg07380026 TERT S_Shore TSS1500 18.0107 9.53E−05 0.652038674 0.740083253 cg07382347 RNF39 Island Body 6.98191 2.34E−06 0.406932169 0.5622676 cg07382943 IGLON5 Island Body 5.36098 0.000192 0.432114708 0.564647343 cg07303092 NKX2-5 S_Shelf TSS1500 9.89379 6.50E−08 0.329076045 0.498830912 cg07407736 MGAT4D Body 4.21075 0.003683 0.281324196 0.350332998 cg07411620 EVX2 N_Shelf TSS200 5.35624 1.24E−05 0.358021001 0.47450737 cg07416664 GALR3 Island 3UTR 5.60422 0.001283 0.618287264 0.694581626 cg07434271 PAX6 S_Shore TSS1500 21.5421 6.22E−16 0.144036724 0.348245551 cg07440414 TCF7 Island TSS1500 17.4859 4.22E−11 0.158247908 0.317328809 cg07440775 GAD2 Island Body 8.64491 2.01E−05 0.38701995 0.502670929 cg07447773 AKR7A3 N_Shore Body 8.68543 0.005234 0.691376868 0.75700303 cg07483304 HOXA10 S_Shelf Body 3.70733 0.008254 0.372224738 0.448420414 cg07502389 NEFM Island TSS1500 15.5495 2.62E−08 0.281389257 0.408542561 cg07515422 SPT8N4 N_Shelf Body 24.6972 6.61E−09 0.514467633 0.716035075 cg07516252 RECS Island TSS200 21.5435 8.49E−13 0.271671274 0.488728262 cg07522913 HOXA3 S_Shelf 5URT 9.74608 0.000221 0.511928783 0.057686315 cg07529210 KDM2B S_Shore TSS1500 5.02938 0.002588 0.354769687 0.445178188 cg07536910 EVX2 N_Shelf 1stExon 8.96009 1.74E−07 0.263254118 0.416078784 cg07539795 PCDHGA12 Island 1stExon 17.5279 4.24E−06 0.595596938 0.719313131 cg07541200 LOC648987 S_Shore TSS1500 4.22398 0.007637 0.325952015 0.407391053 cg07545037 HOXC6 S_Shore 3UTR 517.823 4.58E−08 0.711818597 0.826766354 cg07552803 NEFM Island TSS1500 18.6549 8.12E−10 0.169502645 0.291078075 cg07578695 SLC34A2 Island TSS1500 6.6203 7.80E−06 0.151460871 0.267796383 cg07592963 KCNQ1DN Island TSS1500 6.22493 1.62E−07 0.178661897 0.345640134 cg07595776 LOC101929154 Island TSS200 9.45963 6.84E−10 0.135370743 0.28342468 cg07606384 CYP26C1 Island Body 6.63471 0.000374 0.441954225 0.533425283 cg07615087 SIM2 N_Shelf Body 3.58602 0.0075 0.234275619 0.30896857 cg07653946 GREB1L Island 5URT 10.8697 4.18E−08 0.259665205 0.411714958 cg07660671 SOX7 S_Shore Body 15.7414 3.72E−08 0.431462163 0.582563717 cg07696699 FAM1843 Island TSS1500 5.7036 5.98E−07 0.210884331 0.39049271 cg07702750 TRIL Island TSS200 9.16408 8.11E−08 0.128885084 0.271552312 cg07703462 SSPO Island Body 13.3702 8.23E−13 0.266026585 0.580334178 cg07778029 HOXA9 Island 1stExon 5.61277 0.000322 0.186206651 0.280611154 cg07785447 GSC Island Body 9.13622 2.33E−08 0.154242667 0.304139342 cg07788369 DLX6-AS1 Body 18.7668 3.09E−09 0.181249002 0.294584467 cg07791578 KAAG1 Island 1stExon 11.9854 5.46E−10 0.171653952 0.312987076 cg07793808 KDM28 S_Shore TSS1500 4.66737 0.000325 0.153938716 0.268187382 cg07802350 HOXD13 Island TSS1500 8.17406 5.10E−10 0.177210008 0.36701786 cg07838106 AKR7A3 Island TSS200 49.5186 1.35E−08 0.472668295 0.602705594 cg07855083 SATB2 Island 5URT 3.0641 0.003704 0.13304088 0.225666628 cg07860213 PRDM14 Island Body 12.7997 1.90E−12 0.228971578 0.474873856 cg07871947 SLC12A5 Island TSS200 6.62281 0.000838 0.171775677 0.0249920988 cg07877559 CTNND2 N_Shore Body 6.66452 7.54E−05 0.202223943 0.30331834 cg07879739 GALR3 Island Body 9.06369 1.10E−08 0.174734818 0.336442663 cg07881405 IRX1 Island Body 9.2586 7.53E−10 0.110754914 0.261654085 cg07938743 OTX1 Island Body 5.73923 0.003255 0.631908295 0.702495333 cg07939856 EVX2 Island TSS1500 7.79595 3.58E−05 0.093124488 0.177740516 cg07974511 OTX1 Island Body 10.2 5.33E−11 0.310228367 0.558914471 cg07991112 HIST1HB1 S_Shore TSS1500 13.615 1.05E−12 0.124471345 0.295487683 cg07995594 RUNX3 Island TSS1500 3.2751 0.003527 0.569073647 0.659312249 cg08022244 PTPRN2 Island Body 3.64082 0.000698 0.134820109 0.23204121 cg08034867 ABR Body 12.7754 0.000397 0.679664279 0.765799727 cg08066129 SIM2 N_Shelf Body 8.82211 3.91E−07 0.291328891 0.444891606 cg08079908 HOXD8 S_Shore 3UTR 8.0269 7.71E−07 0.333793427 0.480905144 cg08085357 SOX2-OT N_Shore Body 30.8979 7.23E−07 0.255685122 0.338520537 cg08089301 HOXB4 Island 1stExon 7.18954 4.74E−07 0.114534278 0.256682763 cg08116462 PAX6 N_Shelf 5URT 10.8795 1.31E−08 0.225803489 0.381389378 cg08118159 SPTBN4 Island Body 3.49625 0.00305 0.34717212 0.435882779 cg08126211 KAAG1 Island 5URT 16.5502 4.70E−10 0.29338311 0.452258586 cg08138435 BATF3 S_Shore TSS1500 7.03017 2.21E−07 0.200252405 0.377102944 cg08163199 HLA-1 Island Body 5.49491 5.01E−05 0.100546598 0.199758134 cg08182446 GABRG3 Island TSS1500 21.2981 1.02E−12 0.130479117 0.313677633 cg08196968 PPP1R14A Island Body 4.97821 2.24E−05 0.418621778 0.570853853 cg08203284 HOXA4 S_Shore TSS1500 12.7514 5.72E−06 0.60464538 0.720686343 cg08206318 PITX1 Island 3UTR 15.014 2.56E−11 0.272170181 0.449239288 cg08227526 LRAT N_Shore 5URT 8.03662 1.12E−05 0.182168624 0.267664288 cg08231710 MMP23B Island TSS1500 10.1429 1.85E−09 0.384275077 0.614727335 cg08247587 TTC6 N_Shore Body 11.5921 1.72E−09 0.209435862 0.374884737 cg08274589 SHF S_Shore 5URT 110.22 1.58E−09 0.447071526 0.586194528 cg08274887 POU4F1 N_Shelf TSS200 12.474 1.30E−07 0.171151833 0.275588763 cg08279075 IRX3 Island Body 3.4654 0.00012 0.18910068 0.339620705 cg08305551 MAST1 Island Body 9.40142 6.52E−07 0.101244984 0.219732621 cg08307030 ISL2 Island 3UTR 6.02195 0.000557 0.196708126 0.258678975 cg08309809 TNFRSF10C Island 5URT 20.485 1.48E−11 0.274621861 0.494917212 cg08318726 TBXS N_Shore 5URT 6.32419 0.000935 0.117928388 0.179148543 cg08325845 HLA-J Island Body 5.13258 3.62E−05 0.134642485 0.247463407 cg08370178 LOC648987 N_Shelf Body 6.81612 0.00697 0.436645767 0.50275998 cg08373003 PITX1 S_Shore Body 14.9597 8.89E−07 0.252567718 0.372098099 cg08381274 GDF6 N_Shelf Body 15.4612 7.01E−08 0.492104844 0.665550433 cg08481536 PAX3 Island Body 12.7624 1.03E−12 0.173996353 0.390700297 cg08457898 EPHX3 S_Shore 5URT 22.7283 1.71E−11 0.204066523 0.344169793 cg08465346 HOXC4 Island 5URT 3.81433 0.001851 0.483110791 0.577335322 cg08473380 OTX1 Island Body 22.5708 5.88E−10 0.364137053 0.513038626 cg08479590 HOXA11-AS S_Shore TSS1500 5.00923 0.000587 0.167978388 0.244806283 cg08480826 LINC00461 Island TSS1500 7.053232 0.000229 0.121449564 0.200037248 cg08481464 C10orf11 N_Shore Body 29.2066 9.33E−05 0.746482543 0.827638813 cg08481491 ALDH1L1 S_Shore TSS1500 6.29713 0.009152 0.658953767 0.720887737 cg08492173 IRX1 Island Body 4.53866 0.000106 0.28864169 0.402412636 cg08499046 PAX6 N_Shore 5URT 13.6643 1.56E−11 0.283545371 0.516088816 cg08509840 SLC41AS TSS1500 7.52402 0.000205 0.412567802 0.491769202 cg08511651 HIST1H1A N_Shelf 1stExon 6.76149 1.84E−05 0.477757163 0.610183281 cg08547691 NTN1 Island 3UTR 4.67394 8.23E−05 0.139178866 0.256739873 cg08555653 LINC01391 S_Shelf Body 6.86616 2.55E−05 0.34641655 0.452882498 cg08575330 SP9 Island 1stExon 15.0931 1.87E−08 0.067789705 0.154136625 cg08557845 NEUROG1 N_Shore 3UTR 8.01439 0.00019 0.271920848 0.354515533 cg08598654 DLX6-AS1 Island Body 4.63629 0.000894 0.399000682 0.494693392 cg08599448 ACP5 Island 5URT 3.81295 7.14E−08 0.165191888 0.297231146 cg08613144 LYPD1 Island 5URT 2.97882 0.005611 0.435526518 0.525428249 cg08622757 MNX1 Island Body 4.36751 0.002423 0.189796327 0.262379154 cg08657492 HOXA4 S_Shore TSS1500 7.25973 0.000893 0.38263386 0.450298616 cg08680048 ITPKA Island Body 11.8852 1.23E−11 0.212373575 0.432674401 cg08694699 ESRP2 Island TSS200 4.42589 0.001632 0.389096586 0.484984749 cg08702941 BEND4 S_Shore TSS1500 10.107 0.001991 0.782138853 0.846029111 cg08703872 C2CO4D Island 5URT 5.68938 0.000134 0.608515842 0.741496891 cg08704562 SPRED3 S_Shore Body 6.33441 0.001281 0.443122868 0.529453613 cg08812054 HOXC4 S_Shore TSS1500 6.64908 0.000915 0.26592445 0.327364006 cg08726248 DRD4 Island TSS1500 35.9285 3.66E−11 0.20304968 0.354255703 cg08745288 DLX6 S_Shelf Body 5.3534 0.007822 0.554399248 0.621538333 cg08746900 IRX1 Island Body 7.98462 1.32E−07 0.235090226 0.376738594 cg08748969 CPM S_Shore TSS1500 5.21554 0.002184 0.542231605 0.61301798 cg08781140 MYO15B Island 1stExon 9.68095 5.47E−13 0.15131809 0.404888232 cg08784129 PAX6 S_Shore Body 15.8728 7.19E−10 0.351740274 0.547383227 cg08787968 WT1 S_Shelf Body 7.20922 4.09E−05 0.498584981 0.620879737 cg08793877 LINC01143 Island Body 9.28811 7.39E−09 0.149568198 0.307441445 cg08813062 SP9 Island TSS1500 12.6472 1.63E−06 0.168808022 0.266017165 cg08825084 EN1 N_Shelf Body 9.86512 3.29E−05 0.207569984 0.27952679 cg08828819 PON3 Island TSS200 5.99266 5.99E−06 0.17246284 0.235199425 cg08829841 FOXI2 Island TSS200 8.19331 0.000157 0.319398318 0.409729222 cg08858649 TRIM15 Island Body 3.20734 0.007901 0.192300553 0.279740069 cg08865099 EVX1 S_Shelf TSS1500 20.1282 1.16E−06 0.493333767 0.656054414 cg08874512 IGLON5 S_Shore Body 10.9424 1.30E−09 0.242252623 0.428837938 cg08879910 HLA-J Island Body 5.89136 1.03E−06 0.175856732 0.349495103 cg08898155 PON3 Island TSS1500 9.00469 1.22E−08 0.257349287 0.417773164 cg08902977 PLLP Island Body 7.75936 9.17E−07 0.167494084 0.300486749 cg08927006 HPSE2 S_Shore Body 6.56103 5.77E−05 0.114677434 0.218402815 cg08938793 HOXA10-AS N_Shore Body 3.71061 0.000525 0.263115246 0.384330657 cg08964780 HOXA10-AS Island Body 4.27115 0.007627 0.094318377 0.150416 cg08965276 LMF1 Island Body 5.48137 0.000238 0.229057464 0.334592099 cg08971771 DLX4 Island Body 5.07662 0.004103 0.321769942 0.385341372 cg08973919 SIM1 Island Body 11.1061 2.49E−08 0.284362996 0.442438221 cg08979895 PITX2 N_Shore Body 13.2288 4.10E−09 0.305618018 0.451086939 cg08985979 LOC90768 S_Shore Body 8.5874 9.30E−05 0.539884202 0.661333212 cg09010671 POU4F1 N_Shelf TSS200 12.4516 2.04E−08 0.153113426 0.259183834 cg09016822 IER3 Island TSS200 8.29857 3.99E−07 0.202852156 0.352054082 cg09041678 PAX6 S_Shore TSS1500 27.2427 4.88E−09 0.280274694 0.41761798 cg09042277 TBX5 Island 5URT 8.77965 2.57E−06 0.201589709 0.312155434 cg09046055 PRR18 Island 1stExon 17.4382 1.20E−07 0.254259659 0.357762707 cg09048251 LOC201929154 Island TSS200 7.64012 3.67E−09 0.086145844 0.240297117 cg09050331 PRDM14 S_Shore TSS1500 17.9013 5.79E−05 0.547084194 0.646639717 cg09064304 MGAT4D Island Body 6.26489 3.73E−05 0.087816312 0.198124567 cg09071155 EVX1 N_Shelf TSS1500 5.09354 0.000252 0.246011805 0.345445089 cg09093485 RNF39 Island 3UTR 6.93844 0.006502 0.721870713 0.775979485 cg09098195 BEND4 S_Shore TSS1500 5.66929 3.02E−05 0.418619865 0.554878 cg09105193 CPM Island TSS1500 16.4916 9.13E−07 0.205222605 0.315148295 cg09127400 IER3 Island TSS200 7.27736 1.71E−06 0.197831081 0.341745486 cg09133092 DRD4 S_Shelf Body 10.0972 6.75E−05 0.434975977 0.542118756 cg09135656 PCDHB15 N_Shore 1stExon 6.73573 6.78E−05 0.598642101 0.718853374 cg09143801 SEMA6C Island Body 3.60127 0.001319 0.657554874 0.752062278 cg09156097 PRO51 Island 1stExon 15.1972 4.05E−10 0.294551667 0.494254824 cg09164580 GDF6 Island Body 5.38258 0.000306 0.261103078 0.371037677 cg09180848 BNC1 Island TSS1500 6.29981 0.00013 0.330962189 0.436571435 cg09185099 HOTTIP S_Shore Body 7.55625 0.000148 0.264218328 0.350944839 cg09194159 HOXB4 Island 1stExon 6.20174 0.000106 0.13279474 0.229663663 cg09224689 PAX6 N_Shore 5URT 12.5203 2.34E−15 0.216375987 0.509931735 cg09229918 SYNGR3 Island Body 8.11522 2.21E−08 0.263455755 0.441419542 cg09233651 TBX5 Island Body 4.9558 6.34E−05 0.285938039 0.420097379 cg09234518 NEFM Island TSS1500 12.6959 5.11E−09 0.104511177 0.237903817 cg09234616 WT1 Island TSS1500 7.97392 1.38E−08 0.186676133 0.346339232 cg09236434 KIAA1211L Island TSS1500 8.66357 6.40E−08 0.289038912 0.464051653 cg09238180 HGXD8 Island TSS1500 11.5125 9.01E−08 0.075343361 0.173472059 cg09252999 DKFZp686K1684 N_Shore TSS1500 8.51775 4.21E−08 0.112451065 0.245507505 cg09257796 GABRG3 Island TSS1500 5.51368 0.000112 0.307840527 0.488485 cg09257824 WT1 N_Shelf Body 9.47964 4.56E−08 0.155637375 0.288150506 cg09279736 RNF39 Island Body 9.16597 4.96E−05 0.487407619 0.606497329 cg09299055 ITPKA Island Body 15.9638 1.22E−07 0.445718078 0.595180545 cg09523705 HOXB-AS1 S_Shore Body 5.64094 0.000228 0.321222067 0.429902202 cg09359114 DLX5 S_Shelf Body 5.40576 1.85E−05 0.326755574 0.451554265 cg09982096 PAX6 S_Shore Body 12.9358 4.94E−10 0.148697237 0.309017816 cg09411999 HCXA10 Island TSS1500 3.75679 0.006207 0.366743341 0.450001444 cg09491991 ISM1 Island TSS1500 8.09259 2.14E−08 0.131391309 0.292686298 cg09498572 LINC000403 Island Body 6.2762 4.74E−06 0.199176674 0.328912662 cg09500443 SCG33A1 Island 1stExon 9.96248 2.04E−09 0.170769476 0.351729518 cg09502866 KCNQ1DN Island TSS1500 9.57298 0.00014 0.464518085 0.541069515 cg09509183 IRF6 Island TSS200 13.907 5.46E−08 0.140147101 0.267438327 cg09515953 PPP1R14A Island TSS200 30.9543 1.78E−08 0.504652565 0.651268467 cg09537620 PAX6 Island TSS1500 12.8597 1.91E−13 0.212198392 0.473850545 cg09554509 HOTTIP Island TSS1500 5.64974 0.000439 0.323328364 0.423086768 cg09554951 KCNQ1DN Island TSS1500 17.7183 1.58E−09 0.279655182 0.412002141 cg09585183 TLX1 N_Shelf TSS1500 7.88302 0.000194 0.41390511 0.509769172 cg09591524 HOXA3 Island Body 6.04615 0.000798 0.515768388 0.592310485 cg09595479 PRPH Island 1stExon 4.03112 0.004014 0.375457126 0.451033251 cg09614415 FOX12 Island TSS200 13.8358 3.85E−09 0.246993008 0.414592934 cg09622982 LYPD1 Island 5URT 3.93256 0.002853 0.205485304 0.288362735 cg09639151 PCDHGA12 Island 1stExon 9.28526 1.52E−08 0.303918526 0.510654158 cg09656389 PAX6 Island TSS1500 11.8739 8.85E−11 0.213471005 0.386318653 cg09661370 HOXA11-AS Island Body 12.4182 3.76E−05 0.323708512 0.39229582 cg09671258 LHX4 Island Body 8.45738 5.15E−11 0.115852118 0.288785625 cg09672187 IRX4 Island 5URT 5.64327 0.000101 0.367022354 0.493542719 cg09672383 POMC N_Shore 5URT 13.6288 1.19E−05 0.481664279 0.600165273 cg09686443 CCDC37 Island TSS200 9.79307 1.88E−07 0.104062729 0.221003128 cg09695430 WT1 N_Shore Body 4.43415 0.005337 0.321396093 0.586171233 cg09696159 LINC01391 S_Shore Body 4.2084 0.004215 0.417994752 0.492784343 cg09704116 HOXB-AS3 N_Shelf TSS1500 6.37315 0.000436 0.221848094 0.293098957 cg09706122 ATP5G2 Island TSS1500 3.48179 0.002185 0.174490622 0.278877416 cg09709472 SOX7 S_Shore Body 12.612 1.14E−05 0.41586096 0.520380899 cg09751996 IRX1 Island Body 12.0422 4.61E−11 0.29624417 0.497267249 cg09767822 DBX1 Island Body 26.5295 7.96E−13 0.188538735 0.35250637 cg09792881 DMRTA2 Island Body 12.5702 3.85E−07 0.090862984 0.174475151 cg09797337 LINC00403 S_Shore Body 11.745 2.46E−09 0.102274235 0.251774041 cg09803262 DLX6-AS1 Body 9.16754 2.97E−07 0.243288226 0.357295923 cg09826050 EDRRG N_Shore 5URT 7.92817 6.64E−06 0.509534119 0.648659923 cg09854626 LMF1 Island Body 12.658 2.83E−07 0.191658395 0.350935962 cg09857513 WNT6 N_Shore Body 9.36084 1.16E−05 0.438390302 0.568246424 cg09866983 PRAC2 S_Shelf TSS1500 5.39714 4.69E−05 0.196192305 0.313547852 cg09868598 LOC642366 Body 6.27552 0.000105 0.223075509 0.309642361 cg09888562 IRX3 S_Shelf Body 4.1908 0.001498 0.116633047 0.205114528 cg09955588 GFI1 Island Body 4.33866 0.00487 0.423493545 0.494465208 cg09935824 HOXA-AS3 S_Shelf Body 7.98742 8.15E−05 0.148676625 0.230635268 cg09938227 C1orf61 N_Shore 5URT 7.66101 6.61E−06 0.395072548 0.54391396 cg09983051 MAL Island TSS200 4.59513 0.002886 0.174059019 0.243575552 cg10004780 ARHGEF4 S_Shore Body 9.76679 1.26E−06 0.370631819 0.504917566 cg10035294 PAX3 N_Shelf TSS1500 5.33268 9.60E−05 0.120569757 0.226497141 cg10060574 IER3 Island TSS200 9.04605 1.85E−05 0.221470546 0.330330437 cg10068417 DLEU1 N_Shore Body 15.9542 2.83E−07 0.584037612 0.674070495 cg10074409 IRF6 Island 1stExon 20.5702 7.59E−13 0.22023883 0.334189096 cg10084644 GPC2 Island TSS1500 24.2026 2.84E−10 0.136373097 0.25506711 cg10094489 SHH Island Body 17.2643 1.00E−10 0.276750212 0.468147141 cg10114327 QRFPR Island 1stExon 8.3273 9.10E−05 0.217852651 0.29205139 cg10122865 OTX1 Island Body 4.17075 6.58E−05 0.353863274 0.495327084 cg10131194 GDF6 Island Body 6.57385 0.000186 0.149709165 0.236587793 cg10137231 REC8 N_Shore TSS1500 56.1001 2.94E−11 0.468514256 0.620307525 cg10146929 HIST1H1A N_Shelf 1stExon 6.20351 5.75E−06 0.391991233 0.538500289 cg10147797 SDR42E1 Island 1stExon 9.04143 6.35E−10 0.228458416 0.445737993 cg10159241 APC2 Island Body 5.01963 0.000348 0.383859226 0.495420326 cg10172669 BEND4 Island 5URT 12.2644 7.66E−10 0.204238498 0.358497387 cg10195901 HORMAD2 Island TSS200 10.0352 7.47E−07 0.323685537 0.453139761 cg10242160 ZAR1 Island Body 10.7264 3.07E−07 0.318371089 0.433800141 cg10242602 ZFP42 Island TSS200 11.1904 2.05E−08 0.337679466 0.517529672 cg10245273 SOX1 S_Shelf 3UTR 4.67187 0.000924 0.242752784 0.324129913 cg10245988 SYN2 Island ExonBnd 9.40017 4.09E−07 0.214975 0.348727117 cg10288510 PROX1-AS1 Island Body 10.8597 8.31E−07 0.239522732 0.351650141 cg10300188 KCNQ1DN Island TSS1500 11.5991 0.000331 0.464158333 0.537704162 cg10303487 DPYS Island 1stExon 6.041 6.26E−06 0.062408253 0.167982564 cg10331779 CTNND2 S_Shore TSS1500 48.0325 3.55E−10 0.177938553 0.272077057 cg10334767 ZNF518B Island TSS200 10.5729 6.73E−08 0.179857128 0.303992066 cg10344832 SKOR1 Island Body 17.3533 2.39E−09 0.222910407 0.341060832 cg10347335 TACSTD2 Island 1stExon 6.05381 0.000208 0.233464909 0.306597505 cg10356613 LHX1 Island TSS1500 6.94183 6.24E−06 0.081029232 0.175810062 cg10357657 JAK3 Island 5URT 10.2996 1.32E−05 0.157145195 0.257115862 cg10366797 HOXC4 Island 5URT 29.0514 9.48E−07 0.592405349 0.687697859 cg10421979 CHAD Island TSS200 3.92485 0.000329 0.227963574 0.363060729 cg10439765 SLC12A5 Island 5URT 15.1335 1.07E−09 0.095726603 0.200327056 cg10456990 SIM2 N_Shelf Body 3.71941 0.009115 0.198960606 0.265556813 cg10475970 PCDHGA8 Island 1stExon 33.2528 4.66E−12 0.329747031 0.482509091 cg10479082 TRIL Island Body 16.1443 1.75E−11 0.171703895 0.337100052 cg10500362 SIM2 S_Shore Body 16.8724 6.02E−06 0.529984031 0.635513283 cg10505993 SCG33A1 Island TSS200 10.7966 5.93E−07 0.196189595 0.302974744 cg10511988 VAX1 N_Shore 3UTR 5.62292 8.34E−05 0.269122054 0.407997249 cg10526374 ASCL2 Island TSS1500 11.4498 8.70E−08 0.05920574 0.168102887 cg10532489 SEZ6L2 Island Body 5.81626 4.20E−05 0.331679242 0.474720293 cg10536898 LOC642366 Body 20.1531 1.07E−11 0.358995713 0.536003475 cg10537450 NBPF8 Island Body 22.1861 8.67E−06 0.438182052 0.506966818 cg10588696 ISL2 S_Shelf Body 5.76378 5.41E−05 0.35049756 0.470118863 cg10588962 HOXB-AS3 N_Shore TSS200 7.06082 9.94E−08 0.320369589 0.486445386 cg10603004 NKX2-5 Island TSS200 9.7666 2.77E−06 0.180637448 0.288563559 cg10657141 HOXA11-AS N_Shore Body 16.2069 1.70E−06 0.276586925 0.372236899 cg10659805 DLXG-AS1 N_Shelf Body 8.90602 4.90E−08 0.243029012 0.393836448 cg10660256 BHMT 5URT 49.5233 5.34E−11 0.484227194 0.639939434 cg10682560 DKFZp686K1664 N_Shore Body 12.4474 1.14E−06 0.480962716 0.620314452 cg10689404 LHX1 Island 3UTR 11.2255 1.86E−06 0.242936302 0.355552655 cg10689528 GDF6 Island Body 7.4001 1.39E−06 0.134920426 0.250564504 cg20724867 HOXA10-HOXA9 N_Shore Body 17.0683 1.04E−06 0.58984669 0.695939616 cg10741025 PAX9 Island Body 3.78791 0.000821 0.129727883 0.215902894 cg10745499 SIM1 Island Body 6.33913 0.004526 0.350666025 0.429264566 cg10757144 PCDHB15 S_Shelf 1stExon 4.25491 0.000808 0.177543849 0.267551624 cg10767223 TERT Island TSS1500 18.8804 4.84E−10 0.379046877 0.526890232 cg10782668 PDX1 Island Body 5.30538 0.001217 0.140135129 0.205854312 cg10798171 ICA1 S_Shore TSS1500 7.14828 2.41E−06 0.201468657 0.350483933 cg10798682 SLC41A3 1stExon 36.4821 2.41E−07 0.607802984 0.730013788 cg10823473 ISL2 Island Body 15.6547 2.23E−08 0.22424549 0.353090456 cg10835584 ONECUT2 Island Body 10.3083 4.74E−07 0.057105257 0.144544992 cg10850930 SIM2 N_Shore Body 6.86049 0.002208 0.613732798 0.687522869 cg10859133 OTX1 Island Body 25.6545 4.87E−09 0.410904984 0.537526182 cg10865856 RNF39 Island 1stExon 4.97877 0.000231 0.092297555 0.178510471 cg10895452 EN1 N_Shelf Body 5.58778 5.78E−05 0.209537907 0.307907708 cg10911990 PAX9 S_Shore 5URT 3.28841 0.008825 0.243159468 0.324198692 cg10916998 FLOT1 Island TSS1500 9.14504 3.44E−08 0.338997498 0.530957659 cg10930308 RNF39 Island Body 5.74199 2.90E−05 0.459534134 0.602776067 cg10938374 IER3 Island Body 10.6163 1.18E−11 0.150986656 0.377955083 cg10940462 NRXN2 Island Body 3.8876 0.000247 0.250057522 0.387689454 cg10945917 C1orf61 TSS1500 4.11519 0.000722 0.329809047 0.447263753 cg10953317 CD300A TSS200 12.563 3.49E−05 0.371705338 0.471515052 cg10954251 TBR1 S_Shore TSS1500 5.1599 0.000199 0.147559542 0.245399118 cg10954469 LINC01143 Island Body 9.36353 1.41E−07 0.134322719 0.2443784 cg10957151 HOXD9 Island TSS1500 4.61975 0.003628 0.281270691 0.345664464 cg10964421 TNFRSF10D Island Body 11.951 1.78E−09 0.132322652 0.279332034 cg10977770 ESRRG N_Shore 5URT 13.182 5.66E−09 0.286516066 0.429441477 cg10979436 HOTTIP N_Shelf Body 5.04735 0.000134 0.095966249 0.188729395 cg10983544 TRIM58 Island TSS200 16.2842 5.57E−09 0.114201938 0.244805359 cg10989897 ELAVL4 Island TSS200 14.845 2.24E−10 0.108558528 0.224069455 cg10997718 TBR1 S_Shore Body 12.5008 6.11E−09 0.273866972 0.431588697 cg11015251 HOXA4 Island TSS200 14.2562 1.24E−07 0.335861334 0.44644321 cg11022452 ALDHIL1 S_Shore 1stExon 5.22667 1.07E−05 0.145257533 0.296916145 cg11025705 KIFC2 Island Body 86.1394 2.95E−12 0.41849231 0.587120737 cg11060441 SSPO N_Shore Body 35.0177 1.07E−06 0.654528372 0.774756596 cg11075029 SHF S_Shore 5URT 8.39843 1.37E−07 0.187656884 0.34894378 cg11076994 TCF15 Island TSS1500 20.3299 1.32E−08 0.139085556 0.234768108 cg11080549 NR2F1-AS1 S_Shore Body 10.7771 2.27E−06 0.41834253 0.582399451 cg11086764 NR2E1 Island Body 9.22958 4.32E−06 0.127293425 0.212657107 cg11097541 HOXA10-AS N_Shore TSS1500 11.2992 4.96E−06 0.244468791 0.333403646 cg11101117 LBX-AS1 S_Shore Body 7.64953 1.77E−07 0.275642303 0.466333029 cg11111460 LOC90768 Island Body 11.2971 1.98E−05 0.574232802 0.711423401 cg11128216 PAX6 Island TSS200 11.5677 8.08E−10 0.350681107 0.554415187 cg11162118 DKFZp686K1684 Island TSS1500 6.58767 1.74E−07 0.335928761 0.510457651 cg11168433 KCNQ4 Island ExonBnd 32.7561 4.98E−08 0.567723891 0.711173929 cg11179081 HIST1H25K S_Shore 3UTR 3.89317 0.004639 0.494047639 0.582601063 cg11196848 FGF20 S_Shore TSS200 18.5709 0.00017 0.700089395 0.788481859 cg11207534 ANK1 Island Body 12.9428 2.45E−05 0.147028214 0.22199097 cg11229513 PRDM14 Island TSS1500 8.03748 2.91E−05 0.212457433 0.296093825 cg11229862 CCDC37 Island TSS200 8.35884 3.06E−06 0.138860612 0.247322508 cg11233384 AKR7A3 S_Shore TSS1500 8.6911 6.36E−05 0.400233891 0.524285929 cg11280525 LINC00461 Island Body 20.6051 1.66E−08 0.265839864 0.394563705 cg11282676 HISTIH1A N_Shelf 5URT 9.90773 8.65E−06 0.365720756 0.482940576 cg11301556 SLC12A5 Island TSS200 6.41751 0.00015 0.157213418 0.249876458 cg11320910 GATA4 Island 5URT 29.3702 7.88E−09 0.372212651 0.495148717 cg11325997 AMH Island Body 30.2868 5.85E−10 0.271474938 0.414804552 cg11330740 NKX2-5 Island Body 9.96574 0.000377 0.478069682 0.561559657 cg11335335 DRD4 Island Body 10.1416 0.000131 0.592375979 0.708972825 cg11359133 EVX2 N_Shelf TSS200 7.39901 6.58E−07 0.331291113 0.451562689 cg11368643 PCDH815 S_Shelf 5URT 20.4105 2.74E−09 0.268315226 0.393278391 cg11397370 GRHL2 Island TSS200 20.5187 3.28E−09 0.214270804 0.349032242 cg11418477 SP9 Island TSS200 14.6008 4.47E−07 0.176523971 0.279260216 cg11435506 PON3 Island TSS1500 9.91863 3.12E−09 0.307274088 0.489407326 cg11469061 PAX6 Island TSS1500 16.0312 2.80E−14 0.154525969 0.353459059 cg11469778 ARHGEF4 S_Shore TSS1500 4.95756 0.003171 0.465942168 0.54295198 cg11471772 SIM1 N_Shelf TSS200 4.85052 5.35E−05 0.175868906 0.292676685 cg11472521 DLX4 Island Body 4.2666 0.001663 0.431103093 0.531439207 cg11475550 NPY Island 5URT 3.43701 0.008535 0.196253386 0.257823266 cg11485451 TCF7 Island TSS200 10.8092 8.03E−10 0.309068173 0.52311637 cg11485595 NPR3 Island Body 8.48464 1.01E−07 0.112023276 0.24435177 cg11500797 DLX5 Island Body 5.39912 0.000378 0.238454151 0.32557966 cg11508828 SSPO Island Body 13.3887 2.12E−05 0.49084586 0.595345939 cg11523020 UPK3A S_Shore Body 22.411 9.05E−08 0.382621393 0.507177212 cg11529250 LIPK3A N_Shore TSS1500 5.30122 4.82E−05 0.140324321 0.243532794 cg11538641 SIM1 N_Shelf Body 4.98232 4.40E−05 0.292735853 0.412089714 cg11540692 SIM1 Island TSS1500 7.40857 8.76E−05 0.248331369 0.323636328 cg11605855 IRX1 Island Body 8.95968 1.98E−08 0.294331442 0.469414206 cg11616651 AMH Island Body 11.3565 1.60E−10 0.21201655 0.423223295 cg11625005 TERT Island TSS1500 4.72634 0.000161 0.374007884 0.490540023 cg11633298 ZFP42 S_Shore 5URT 6.28575 0.000573 0.39812143 0.490456717 cg11644479 ASCL2 Island TSS1500 5.29529 0.005569 0.604096358 0.86798104 cg11648594 MMP23B Island TSS1500 6.11 2.27E−07 0.266911991 0.456886426 cg11651285 NKX2-2 Island TSS200 10.1528 1.59E−05 0.280996806 0.375315826 cg11663667 LRAT S_Shore 5URT 6.74162 2.98E−05 0.436425109 0.579243388 cg11667020 NKX2-4 Island 1stExon 10.353 3.17E−10 0.123798491 0.279686414 cg11667086 DKFZp6S6K1684 Island Body 5.68763 0.0005 0.094874515 0.172254134 cg11671598 PITX2 S_Shelf 1stExon 8.29026 6.90E−05 0.518653171 0.597968606 cg11679455 GATA3 Island Body 6.6013 0.006322 0.798564597 0.858473778 cg11700800 NR2F1-AS1 Island Body 5.23124 0.00204 0.814014127 0.396130636 cg11724134 RNF39 S_Shore TSS200 8.17163 5.71E−06 0.18787539 0.293942616 cg11741189 DKFZp686K1684 N_Shelf Body 10.8418 1.22E−08 0.393777786 0.586224771 cg11754318 HOXD12 S_Shelf TSS200 11.5034 5.45E−07 0.367504876 0.487584232 cg11770080 BHMT Body 4.91959 8.71E−06 0.22203796 0.407015969 cg11772171 ASCL2 Island TSS1500 7.59457 2.00E−06 0.125973509 0.235435424 cg11789612 ITPKA Island Body 11.6506 1.86E−13 0.126004371 0.349646136 cg11800620 ESRRG S_Shore TSS1500 4.98139 2.44E−05 0.300581095 0.432389181 cg11806672 POU4F1 Island Body 23.8502 2.27E−11 0.197183953 0.329205928 cg11823511 BARHL2 Island TSS1500 9.97971 2.33E−09 0.11159678 0.249968451 cg11827910 PAX6 Island 5URT 13.9055 9.73E−10 0.417511434 0.624910141 cg11856320 PDX1 S_Shore TSS1500 10.4005 4.21E−07 0.281451814 0.405942482 cg11853830 NR2F1-AS1 N_Shore Body 24.3189 1.59E−07 0.587821858 0.790941384 cg11854154 AMPD3 Island TSS200 5.02552 0.000692 0.191287998 0.280976332 cg11865196 HNF15 Body 6.64005 0.000264 0.244260691 0.340077945 cg11891393 SIM1 N_Shelf TSS200 9.2349 1.88E−06 0.204055279 0.31220807 cg11904906 LOC151174 N_Shore Body 7.6273 7.44E−06 0.284219535 0.414816179 cg11910375 HOXA-AS3 S_Shore Body 8.37918 0.000237 0.159098216 0.219262914 cg11915729 SCG88A1 Island TSS200 10.1315 2.08E−10 0.119576398 0.310519186 cg11930477 DKFZp586K1684 S_Shelf Body 15.1845 1.86E−09 0.159756347 0.274739177 cg11931731 KATNAL2 Island 5URT 6.84976 8.22E−09 0.172180564 0.360768351 cg11945824 PRRT1 Island 3UTR 6.28934 0.000315 0.601914349 0.672759303 cg11946503 NEUROG1 N_Shore 3UTR 5.33819 0.000487 0.30599935 0.410681973 cg11947493 TNFRSF10D Island 1stExon 10.8915 1.17E−07 0.316887709 0.480101844 cg11949335 TTC22 Island Body 9.29929 0.001219 0.669952473 0.723164626 cg11965311 LOC145845 N_Shore TSS200 4.46979 0.001685 0.877009824 0.447727247 cg11969002 IER3 Island 1stExon 6.89253 9.82E−08 0.279592875 0.481414618 cg12006284 WT1 Island Body 9.87543 5.83E−06 0.305079158 0.418529697 cg12018098 SPRED3 Island Body 3.48044 0.002288 0.297320482 0.3862115 cg12064275 RNF219-AS1 Island Body 4.74441 0.00234 0.380819326 0.400964939 cg12065840 COL14A1 S_Shore 5URT 6.39092 8.92E−07 0.395051158 0.574608368 cg12074182 EVX2 N_Shore 1stExon 11.7178 8.98E−05 0.26933362 0.353775192 cg12090740 BCL2 S_Shelf TSS1500 5.27945 0.00061 0.48011662 0.584088038 cg12110087 HOXA-AS3 S_Shelf Body 6.39135 0.000677 0.187065374 0.247584495 cg12118246 CD300A TSS200 6.36681 0.00232 0.553286029 0.611540382 cg12147489 HOXA11-AS S_Shelf Body 6.06786 9.09E−05 0.190078146 0.285841705 cg12163182 TBX2 Island TSS1500 12.6554 1.78E−05 0.483960977 0.585550949 cg12164596 MNX1 Island Body 17.9596 1.30E−06 0.298142558 0.393756798 cg12171183 KATNAL2 N_Shore TSS200 8.60191 2.18E−10 0.114444297 0.296725188 cg12134886 MYO15B Island TSS200 13.5608 1.33E−10 0.301012461 0.493721663 cg12198813 KCNQ1DN Island TSS200 5.78783 0.000204 0.309642026 0.405589714 cg12200038 TTC6 Island Body 11.4 3.49E−12 0.149843411 0.353821761 cg12213062 SPEG Island TSS200 6.12436 7.14E−05 0.231187526 0.358882629 cg12224030 DLX4 S_Shore Body 8.66417 7.45E−05 0.897200455 0.494769683 cg12233379 C2CD4D Island 5URT 8.51442 0.000122 0.464945878 0.577410727 cg12248614 SPTBN4 Island Body 24.5699 7.82E−08 0.515782742 0.691723995 cg12258785 THRB-AS1 Island Body 5.61478 0.001075 0.593633165 0.67060731 cg12266953 GDF7 N_Shelf TSS200 4.41548 0.000449 0.055960468 0.142984981 cg12268637 HOXD12 S_Shelf TSS200 12.5017 4.89E−08 0.299423647 0.419984697 cg12278467 LINC00461 Island Body 14.2163 5.56E−07 0.213344535 0.29564764 cg12305481 HOXA3 S_Shelf 5URT 4.98264 0.001665 0.468909969 0.546158152 cg12308746 DLX1 S_Shore Body 4.84722 0.001535 0.291872257 0.356015548 cg12509653 LOC145845 N_Shelf Body 3.78509 0.00871 0.265807289 0.345250709 cg12331932 CD300A 5URT 3.82618 6.95E−07 0.282764705 0.437189669 cg12338417 TRIM71 Island 1stExon 14.0698 3.35E−09 0.201685908 0.357514314 cg12349884 HIST1H28K N_Shore 5URT 5.70568 1.54E−05 0.328347509 0.458301584 cg12374721 PRAC2 S_Shelf TSS1500 7.78488 9.58E−12 0.189220395 0.449903944 cg12377139 SPAG6 Island TSS200 11.3725 7.04E−08 0.237886291 0.375570137 cg12384286 TMPRSS2 S_Shore TSS1500 5.66982 0.003553 0.426987391 0.498744822 cg12386646 ITPKA Island Body 8.90491 2.18E−07 0.230532981 0.365584251 cg12393318 LINC00461 Island TSS1500 18.2579 4.22E−10 0.133958562 0.249547709 cg12405189 GATA3 Body 6.84378 0.001064 0.671935047 0.756305319 cg12434982 LHX9 Island Body 45.334 4.16E−09 0.489409488 0.611427414 cg12473406 GREB1L Island 5URT 4.54793 2.48E−05 0.180743432 0.323613619 cg12518146 F2RL1 Island Body 7.7045 1.98E−06 0.341290988 0.486734551 cg12520549 TBR1 S_Shore 3UTR 8.69701 2.84E−05 0.18679993 0.288207703 cg12566152 LBX1-AS1 S_Shore Body 15.3291 8.83E−08 0.382129616 0.554121897 cg12588917 HLA-F Island Body 47.3415 3.89E−06 0.571920973 0.651341273 cg12606911 CD8A Island TSS1500 4.57958 0.000168 0.205685651 0.2938799 cg12610471 SPAG6 Island TSS200 22.5988 6.12E−11 0.219480396 0.865855893 cg12614105 NPY Island 5URT 3.757 0.001888 0.191134272 0.271748995 cg12638154 RNF39 Island Body 12.4791 0.000187 0.670818979 0.772809596 cg12688169 ESRRG N_Shore 5URT 11.1901 4.79E−08 0.405921693 0.566064003 cg12648074 PCDHGC4 N_Shore TSS1500 5.09057 3.05E−05 0.29104175 0.438440579 cg12670547 TBX5-AS1 Island Body 8.16219 1.16E−05 0.327880761 0.487967781 cg12673103 PITX1 N_Shelf Body 11.4342 8.64E−10 0.272483705 0.486955393 cg12676702 DLX1 S_Shore 3UTR 4.85295 0.000373 0.349860511 0.432538939 cg12685539 AMPD3 Island TSS1500 3.91597 0.006058 0.504952116 0.598179044 cg12700904 FBLN2 Island 5URT 6.43555 8.55E−06 0.13515415 0.289085841 cg12786877 RNF39 N_Shelf Body 9.75787 0.000194 0.702471574 0.805423625 cg12782180 LEP Island TSS1500 9.40214 1.65E−08 0.346491947 0.527324681 cg12788467 HNF16 Island TSS1500 4.16374 0.001219 0.312264142 0.401452574 cg12798259 PAX5 N_Shelf 5URT 62.4417 3.06E−08 0.634593938 0.773873091 cg12807794 TCF15 Island TSS200 13.0461 1.61E−06 0.135929877 0.248012886 cg12810523 HOXA-AS3 Island Body 8.95793 1.07E−05 0.121947364 0.201956428 cg12821804 KCNQ1DN Island TSS1500 6.47471 5.65E−07 0.317049374 0.458569091 cg12876356 GFI1 Island Body 7.4366 4.33E−05 0.300802698 0.88211982 cg12883523 SOX17 Island Body 9.11479 8.32E−05 0.293728913 0.384119535 cg12833980 DLEU1 S_Shelf Body 8.01939 0.000172 0.47203124 0.55021399 cg12902206 DNAH1 5URT 6.51081 0.000144 0.543186877 0.674361414 cg12910906 ARHGEF4 Island TSS1500 6.43703 0.00039 0.232164348 0.314418186 cg12928379 DRD4 Island TSS200 5.45491 1.65E−05 0.078505244 0.187030821 cg12945444 GATA4 Island 5URT 18.8458 5.92E−08 0.15996906 0.248073929 cg12953669 BCL2 S_Shelf TSS1500 9.37243 2.04E−05 0.386564078 0.466781778 cg12967914 RNF89 Island 1stExon 4.47364 3.64E−06 0.132900042 0.283289125 cg12969193 HOXD9 S_Shore Body 6.42672 1.88E−05 0.382825116 0.502787577 cg12973941 NKX3-2 Island 1stExon 18.6876 3.90E−11 0.173925328 0.320769502 cg12982522 WT1 S_Shore Body 6.48468 0.000378 0.32771907 0.400272283 cg12984729 ISL2 Island Body 6.45216 1.26E−05 0.268807585 0.391906917 cg12997720 HOXA11-AS Island Body 7.58896 3.91E−05 0.368146048 0.497772888 cg12999453 TBX2 Island Body 3.09508 0.005125 0.223975444 0.321880888 cg15016408 LYPD1 N_Shore 5URT 19.2203 1.06E−08 0.194973351 0.309842982 cg13028628 OTX2 Island Body 4.93906 0.000139 0.274117076 0.389552712 cg13025668 POMC Island TSS200 7.67935 1.08E−05 0.212290791 0.342869899 cg13029700 PAX6 Island 5URT 10.5581 6.18E−09 0.127799755 0.278899162 cg13046271 TCF7 Island TSS200 24.9628 8.66E−10 0.810896534 0.460638793 cg13073778 GSC Island 3UTR 4.92109 1.28E−05 0.331311017 0.491597126 cg13112154 LBX1-AS1 Island Body 15.0407 4.04E−09 0.19466912 0.345043156 cg13115683 HCG9 N_Shore Body 5.74997 1.67E−05 0.256822455 0.428518198 cg13187376 IER3 Island 1stExon 8.18428 2.60E−06 0.178674205 0.232852882 cg13147822 MECOM Island 1stExon 7.56712 3.91E−05 0.265840778 0.39770754 cg13163521 LTK N_Shore Body 7.84086 0.00017 0.42813884 0.506217347 cg13185426 RNF39 Island Body 5.80524 2.00E−05 0.326688835 0.485006807 cg18193289 PITX2 Island Body 22.8617 5.86E−09 0.242346795 0.344768687 cg13210467 GPC2 Island TSS1500 25.1222 2.68E−07 0.496041186 0.606976192 cg13211683 GDF6 Island Body 5.89754 0.00058 0.089594178 0.167606413 cg18280606 HNF1B Island TSS1500 4.02872 0.003296 0.210870806 0.284155771 cg13245152 PAX6 N_Shelf 5URT 14.9118 8.73E−08 0.49202105 0.656317798 cg13301003 WT1 Island Body 8.2718 6.41E−05 0.475607806 0.589881061 cg18805657 RUNX3 Island TSS1500 8.90981 0.000552 0.494884519 0.559488283 cg13331550 SEMA6C Island 5URT 6.80241 5.44E−06 0.221839788 0.377575976 cg13338157 RNF219-AS1 N_Shore Body 9.04608 8.82E−06 0.141680975 0.225800909 cg13865524 SIM1 N_Shelf Body 6.78962 2.12E−05 0.255778125 0.361656149 cg13368756 CTNND2 Island 1stExon 7.62656 3.92E−08 0.096583321 0.219273855 cg13378628 OTP Island TSS200 8.35047 0.000147 0.122864769 0.199601241 cg13381984 LEP Island 1stExon 11.9997 4.55E−06 0.290130926 0.08741142 cg13401893 RNF39 Island Body 9.36716 0.000293 0.670615192 0.772663716 cg13407335 KCNAB8 N_Shelf TSS200 11.9979 2.56E−06 0.589493791 0.711425879 cg13409449 GATA3 Island 5URT 5.72698 1.10E−05 0.229138767 0.872526068 cg13426079 KCNQ4 Island Body 5.77062 0.000648 0.580478484 0.669677869 cg13454842 GATA4 S_Shore 5URT 15.4966 5.53E−10 0.156796369 0.294757757 cg13448627 TACSTD2 Island 1stExon 7.40151 1.12E−05 0.849093956 0.480215532 cg13445199 ZNF5186 Island TSS200 8.45915 2.21E−07 0.157318902 0.277190093 cg15459498 TOX2 Island 1stExon 6.49721 0.000219 0.090607309 0.176815177 cg13476950 DNAH1 5URT 10.0886 4.22E−05 0.409590799 0.529263172 cg13492692 LHX2 Island Body 11.1808 3.73E−09 0.127132857 0.271664542 cg18502125 NBPF8 Island Body 20.6193 2.85E−08 0.520453463 0.688828657 cg13528935 FAM84B Island 5URT 7.88759 7.72E−07 0.136828803 0.255677428 cg13530938 LOC151174 Island Body 6.38122 8.60E−06 0.297450414 0.450294253 cg18595212 FOXI3 Island TSS1500 6.29034 1.04E−05 0.249519568 0.379507738 cg13540960 WT1 N_Shore Body 9.37513 5.68E−08 0.102176159 0.203781828 cg13552869 SEZ6L2 N_Shore Body 8.84234 0.000167 0.397762271 0.489042505 cg13564825 PPP1R14A Island 1stExon 7.72339 1.06E−10 0.213819879 0.452577083 cg13567542 TRIM15 Island Body 4.55736 0.000962 0.17590263 0.275526075 cg13570101 NKX2-5 Island 1stExon 9.3288 1.05E−05 0.120154288 0.221241881 cg18570972 PAX6 Island TSS200 4.35325 0.00365 0.247366536 0.820463685 cg13596833 PAX6 Island Body 15.7471 1.16E−11 0.114125652 0.273748385 cg15604794 DLX4 Island Body 9.63385 4.36E−05 0.220114995 0.304732658 cg19605398 KCNAB3 Island 5UTR 6.14283 5.75E−05 0.175484885 0.298217873 cg13614181 RGCC Island TSS1500 18.6218 8.61E−06 0.463725287 0.557841444 cg13617591 LOC101929076 Island Body 7.41402 4.59E−06 0.187950929 0.24213276 cg13641903 WT1 Island TSS1500 16.8045 7.98E−08 0.326631802 0.458745279 cg13648914 ORFPR Island TSS200 8.76048 8.83E−07 0.05882172 0.15469307 cg13660279 IRX3 S_Shelf Body 3.78295 0.008062 0.321422494 0.890681475 cg13691247 ATP5G2 Island TSS1500 3.41475 0.00145 0.097549792 0.197930448 cg13692446 LINC00403 Island Body 13.2831 1.13E−13 0.103096408 0.306827221 cg13694575 DKFZp686K1684 Island TSS200 6.81459 0.001594 0.20278612 0.268589346 cg18703049 HOXA10-HOXA9 Island Body 19.7056 9.30E−12 0.102623495 0.249122317 cg13715681 PITX1 S_Shore Body 7.72871 8.55E−06 0.339909099 0.477012534 cg13726459 HOXC4 N_Shelf TSS1500 3.85017 0.001381 0.379902319 0.471530557 cg13735819 JSRP1 Island Body 5.42846 8.42E−05 0.242931318 0.357270722 cg13742526 JSRP1 Island Body 6.15767 4.74E−08 0.2853683 0.500366468 cg13752649 HOXB1 S_Shore TSS1500 7.76946 4.54E−05 0.471201535 0.574165242 cg13762320 ASCL2 Island TSS1500 3.70708 0.00097 0.584702372 0.655773566 cg13764778 CHAD Island TSS1500 5.38447 0.001643 0.312652583 0.895388813 cg13774342 DLEU1 S_Shore Body 5.37973 0.004414 0.738923349 0.790965848 cg13784235 NR2E1 N_Shore Body 14.4471 3.48E−10 0.263442388 0.412421966 cg13799941 DLX6-AS1 S_Shore Body 18.5242 7.92E−07 0.349515519 0.449267414 cg13803976 CD8A Island Body 8.74282 1.01E−05 0.150053105 0.238986572 cg13804182 FLOT1 Island TSS1500 9.15684 6.72E−09 0.316988381 0.537997999 cg13813866 SHISA3 N_Shore TSS1500 5.87144 8.16E−05 0.491128384 0.614594949 cg13826247 HOXC5 S_Shore 3UTR 4.62181 0.000297 0.450008462 0.557003626 cg13870865 TBX20 Island 1stExon 10.058 0.000165 0.271168256 0.331129807 cg13885561 KCNQ1DN Island TSS1500 4.06919 0.005038 0.455452411 0.522336063 cg13885965 IER8 Island TSS200 5.88537 2.10E−06 0.224712908 0.399499856 cg13891220 PITX2 Island Body 5.71795 0.000655 0.292976251 0.373341866 cg13891702 PAX6 N_Shore 5URT 14.3954 3.73E−10 0.216900514 0.388425891 cg13918754 RNF39 Island Body 5.22055 7.85E−06 0.163281703 0.298564109 cg13922021 F2RL1 Island Body 39.1967 3.80E−08 0.472491891 0.589959424 cg18929328 FOXI2 Island 1stExon 11.3167 7.87E−07 0.402985542 0.536186531 cg13965612 MIR199A1 TSS1500 9.28348 0.000215 0.592721512 0.680766919 cg13967702 ESRRG S_Shore TSS1500 6.48984 1.40E−06 0.267616488 0.426707968 cg13974394 IRX4 N_Shelf Body 13.8641 2.17E−06 0.247993919 0.34193616 cg13982098 LINC00461 Island Body 17.3128 9.31E−09 0.88450089 0.470155949 cg13990559 ALDOC TSS200 40.7063 4.68E−10 0.24829618 0.387324537 cg14020315 RNF219-AS1 S_Shore Body 3.5105 0.002612 0.319186698 0.402130192 cg14013695 HOXA5 Island TSS1500 7.95389 8.23E−05 0.410005482 0.529976602 cg14015044 TNFRSF10C Island 5URT 15.8398 1.00E−11 0.11000692 0.278207112 cg14015441 DPYS Island TSS200 9.18155 8.54E−08 0.219530945 0.37461871 cg14029489 PRPH Island 1stExon 4.74702 6.51E−06 0.217821895 0.377856606 cg14034197 GR5B1L Island 5URT 5.60749 5.88E−05 0.189781316 0.34136425 cg14044640 HOXA-AS3 Island Body 12.6773 1.46E−11 0.079247903 0.241340992 cg14069965 KAAG1 Island 5URT 9.40757 1.05E−05 0.318146075 0.447658818 cg14084907 TP78 Island Body 5.19343 8.25E−05 0.311378956 0.434132145 cg14035358 EVX2 N_Shelf TSS1500 5.64153 0.000363 0.350296643 0.441828495 cg14097304 MNX1 Island Body 6.78494 4.46E−06 0.222925273 0.341250906 cg14159342 TP73 S_Shore Body 17.0189 3.24E−07 0.462922054 0.596208354 cg14177914 C1orf61 TSS1500 8.04093 3.67E−06 0.45098763 0.606254106 cg14181940 CD800A Body 8.5887 4.75E−05 0.425904459 0.541955 cg14189571 ZFP42 S_Shore 1stExon 16.7846 1.06E−06 0.565770065 0.717826798 cg14210607 LOC101929076 Island Body 5.2511 0.000374 0.345443798 0.452102564 cg14228238 MECOM Island TSS200 8.4457 1.51E−05 0.268087512 0.395772775 cg14230397 PRAC2 N_Shore TSS1500 11.2353 1.25E−07 0.171094905 0.291555508 cg14239515 SPRED3 Island Body 2.9586 0.008382 0.221098705 0.60374516 cg14239592 ZNF274 Island 5URT 22.7394 1.47E−05 0.695863876 0.779931525 cg14244013 TBX2-AS1 Island Body 43.4861 7.43E−09 0.56676283 0.743882747 cg14301212 HCG9 N_Shore Body 6.46623 0.001447 0.386004459 0.478843467 cg14314744 SIM1 Island Body 7.80909 1.84E−08 0.222133657 0.386512518 cg14317513 SIM2 N_Shore Body 7.51912 0.004101 0.632752938 0.705865949 cg14319235 ADGRL4 Island Body 8.37399 3.50E−07 0.296350767 0.449644854 cg14327531 GATA3 Island 5URT 7.03233 8.83E−05 0.360955997 0.479615796 cg14337027 HOXB13 N_Shore 3UTR 8.32211 4.12E−06 0.409713736 0.539429707 cg14345497 HOXB4 Island 1stExon 8.04459 4.86E−06 0.13726798 0.232549221 cg14353137 TCF24 Island Body 10.0356 3.12E−09 0.083449275 0.224472564 cg14355911 TBX2-AS1 Island Body 13.1296 1.45E−06 0.201063822 0.283530198 cg14375937 CYP26C1 Island Body 6.20711 3.62E−05 0.127658284 0.282282321 cg14385245 BNC1 Island TSS1500 9.02353 2.22E−05 0.325916711 0.414728026 cg14410016 HOXD9 Island TSS1500 15.4815 2.43E−06 0.278506601 0.371385909 cg14416523 TBR1 S_Shore Body 8.82131 3.09E−07 0.230427969 0.357622117 cg14425564 LHX2 Body 5.56984 0.000242 0.296992353 0.401853684 cg14428146 NKX2-6 Island 1stExon 11.3326 1.16E−08 0.109763084 0.230381575 cg14432269 TCF24 N_Shore Body 4.64716 0.000125 0.288184922 0.40335312 cg14432910 SKOR1 Island Body 6.48624 0.000152 0.261442813 0.340986787 cg14439629 PAX6 S_Shore 5URT 14.8568 1.24E−07 0.31751593 0.439166152 cg14443953 DKFZp686K1684 Island TSS1500 8.49247 6.68E−08 0.300990197 0.461779536 cg14443169 WNT16 N_Shore Body 8.65511 7.11E−08 0.159698139 0.290554765 cg14451926 NKX2-6 S_Shelf Body 5.98253 0.004013 0.173293454 0.221275318 cg14458068 PCDHGAS N_Shore 5URT 9.31697 1.05E−06 0.387253436 0.537260828 cg14459130 SP9 Island TSS200 15.2367 3.73E−07 0.185629605 0.300829668 cg14461582 TLX1 N_Shelf Body 5.46293 0.000897 0.493007792 0.574722281 cg14473102 HOXD8 Island TSS200 11.5597 1.07E−09 0.094771569 0.22146963 cg14487292 HNF15 Island TSS1500 4.2018 0.000202 0.218627355 0.316154892 cg14501061 ALDOC TSS200 11.8971 1.49E−06 0.222735935 0.338701517 cg14505195 NKX2-3 S_Shelf Body 9.22168 2.87E−06 0.153132987 0.265345336 cg14509403 HORMAD2 Island TSS200 9.75362 1.34E−09 0.203018926 0.356081221 cg14531663 PRRT1 Island 3UTR 3.03181 0.002766 0.436850205 0.516761708 cg14542583 SPRED3 S_Shore Body 4.6049 0.008916 0.379759144 0.444452606 cg14556683 EPHX3 Island Body 266.124 3.34E−10 0.542474054 0.659224788 cg14557487 TBR1 N_Shelf Body 3.66066 0.004468 0.457305126 0.532436533 cg14559388 APC2 S_Shore Body 4.73699 3.24E−05 0.1719199 0.609330983 cg14598764 SLC41A3 TSS200 13.7772 3.87E−07 0.495980536 0.557217475 cg14604681 OTX1 S_Shelf Body 10.2714 4.64E−05 0.436878349 0.533413081 cg14615784 GPR150 Island TSS1500 5.78238 0.000135 0.115498843 0.201100734 cg14619949 F2RL1 S_Shore Body 27.5045 0.00059 0.743681217 0.817955687 cg14626259 SIM2 N_Shore Body 5.72139 3.90E−05 0.360848806 0.481280356 cg14636534 LINC00461 N_Shore Body 21.2142 2.84E−12 0.34872869 0.538823535 cg14649140 HOXA10 Island TSS1500 5.69525 0.000747 0.117110953 0.174707987 cg14652095 HIST1HIA N_Shelf TSS200 13.8896 5.88E−05 0.558506031 0.769997806 cg14657884 TCF24 Island TSS1500 4.174124 0.004633 0.198247367 0.268053214 cg14668747 ALDOC TSS200 12.9714 8.85E−11 0.125141129 0.294460972 cg14678774 MAST1 Island Body 25.7432 2.08E−08 0.438695077 0.606656859 cg14703002 HIST1HIA N_Shelf TSS200 8.87712 2.30E−07 0.250115524 0.388895254 cg14757311 LMF1 Island Body 14.425 6.57E−05 0.66691986 0.742698889 cg14776998 TRIM71 S_Shelf Body 4.9534 0.008232 0.699798395 0.771137798 cg14780466 GDF7 Island Body 21.1984 9.18E−06 0.58389523 0.721946727 cg14781281 HLA-1 Island Body 5.9254 4.36E−05 0.076101451 0.17472572 cg14817655 ZFP42 Island TSS200 13.2241 6.13E−14 0.207407309 0.463723621 cg14879760 LOC642366 Body 7.17192 5.88E−05 0.160745512 0.25210968 cg14889643 BCL2 Island Body 8.44331 4.40E−05 0.238736396 0.357541533 cg14891209 LOC145845 N_Shelf Body 4.29391 0.000307 0.234486124 0.352183654 cg14891410 WT1-AS N_Shore Body 8.98816 5.28E−06 0.384702219 0.513093637 cg14928902 PRDM14 Island 5URT 10.97 1.91E−05 0.256452086 0.33280978 cg14944647 MIR129-2 Island TSS200 14.6789 9.34E−12 0.46661843 1.180557427 cg14958635 NEUROG1 Island 1stExon 13.1422 1.69E−09 0.214916792 0.356502062 cg14974749 HOXA5 N_Shore Body 5.75881 0.000274 0.500670429 0.575029566 cg14982276 TMPRSS2 Island TSS1500 3.63224 0.001124 0.404273458 0.519408513 cg14989164 LHX4 S_Shore Body 6.64443 5.03E−05 0.189609384 0.232277795 cg14991437 HOXD9 N_Shore TSS200 9.55815 5.93E−05 0.146523925 0.216274712 cg14993283 SIM1 Island Body 19.1015 8.99E−06 0.359395264 0.453991102 cg14996783 TRIL Island Body 7.51146 1.56E−06 0.104599512 0.214741819 cg15006175 KIFC2 Island Body 34.2654 1.21E−09 0.530633681 0.781325113 cg15014975 RUNX3 Island TSS1500 3.34261 0.001193 0.409863922 0.5101684 cg15015639 DES N_Shore TSS200 4.04377 0.000601 0.494292476 0.612912345 cg15017067 FAM1843 Body 20.5351 2.24E−08 0.52672407 0.653316202 cg15084543 ADGRL4 Island 1stExon 6.65983 1.33E−08 0.307247155 0.516300318 cg15105182 QRFPR S_Shore TSS200 9.06698 2.09E−07 0.244594852 0.390081244 cg15126179 RNF219-AS1 Island ExonEnd 7.13865 5.90E−05 0.278927226 0.372169088 cg15133719 PAX3 Island TSS200 5.33657 0.001289 0.274900928 0.358737758 cg15137565 NKX3-2 Island Body 4.31047 0.005279 0.252297543 0.300551774 cg15140703 GPC2 Island TSS1500 8.71914 4.35E−08 0.106306797 0.269288291 cg15196805 HOXA4 S_Shore TSS1500 6.8427 0.000704 0.456529295 0.515956444 cg15209808 HORMAD2 Island TSS200 11.3629 2.80E−10 0.242211992 0.419844515 cg15234492 KDM2B S_Shore TSS1500 4.05281 0.000254 0.247401212 0.954253682 cg15235798 LINC00403 S_Shelf Body 5.99752 1.24E−05 0.240463122 0.361136811 cg15236865 DLX1 N_Shelf TSS1500 8.77426 0.005309 0.366566379 0.44088801 cg15244327 TNFRSF10D S_Shore TSS200 8.05348 4.08E−05 0.466352657 0.606350231 cg15255390 HOXB3 Island 5URT 5.17606 0.002093 0.362904902 0.420692273 cg15267232 GATA3 Island Body 4.2943 9.88E−05 0.37801236 0.526824296 cg15299589 ZAR1 N_Shore TSS1500 11.3782 2.08E−05 0.387109348 0.486298745 cg15315685 WT1 S_Shore TSS1500 9.10918 8.72E−08 0.215972564 0.334855181 cg15325875 TLX1 N_Shelf Body 15.7732 1.55E−08 0.368681812 0.55432886 cg15331352 HLA-F S_Shore Body 14.9686 5.24E−06 0.526207264 0.641224131 cg15377285 SOX27 Island Body 4.51036 0.001532 0.345685631 0.438915236 cg15402095 NR2F1-AS1 N_Shore Body 18.7485 2.85E−08 0.435399508 0.635211902 cg15440688 GSC Island TSS1500 9.91801 1.06E−05 0.484183499 0.616127415 cg15444947 TRIL Island Body 14.2428 1.87E−07 0.100575993 0.190176041 cg15445391 WT1 Island TSS200 27.3954 1.22E−08 0.474104171 0.60534503 cg15460093 ADAM8 Island 1stExon 5.90921 4.22E−05 0.118319191 0.224861282 cg15461889 AMPD3 Island TSS200 3.85418 0.005203 0.13689457 0.206358333 cg15475840 LBX1-AS1 N_Shore Body 3.85036 0.009791 0.583117317 0.684176111 cg15491102 DLEU1 Body 9.57168 1.43E−05 0.498687868 0.635026131 cg15500965 PON3 Island TSS1500 25.7927 7.70E−13 0.2011126 0.345445922 cg15505412 IRX1 Island Body 3.3983 0.006498 0.236276122 0.321745448 cg15506477 LYPD1 N_Shore 5URT 4.57141 0.001185 0.20120695 0.226227349 cg15515258 LINC00403 S_Shore Body 7.89795 2.54E−08 0.220295024 0.382486422 cg15552156 DLX1 S_Shore 3UTR 9.20575 5.55E−07 0.243382412 0.360838604 cg15556502 MIR129-2 Island TSS200 11.8391 4.81E−10 0.050271183 0.160005538 cg15564098 EVX1 N_Shelf TSS1500 6.12901 2.83E−07 0.206339087 0.359701272 cg15572489 PTPRN2 Island Body 6.74166 5.35E−05 0.468346529 0.593494276 cg15590989 LOC90768 Island Body 6.10684 1.44E−05 0.361369917 0.506138635 cg15607672 OTX2 S_Shore TSS200 3.99619 0.00059 0.373912446 0.476411263 cg15611279 LINC00461 N_Shore Body 10.6388 8.21E−07 0.081964312 0.169218818 cg15613420 H2AFY S_Shore TSS1500 6.33881 1.80E−08 0.312646591 0.528206235 cg15634877 SPEG Island TSS200 5.96249 0.000472 0.175682686 0.271603799 cg15645660 TTC22 Island Body 7.25656 0.000111 0.517995589 0.603291717 cg15652723 PITX2 N_Shore 5URT 5.44024 0.000554 0.279235258 0.355374546 cg15653173 LINC00403 Island Body 12.676 4.18E−07 0.149868743 0.244933183 cg15660418 HOXC4 N_Shelf TSS1500 3.11084 0.007086 0.395937178 0.486979967 cg15665400 LYPD1 Island 5URT 3.42973 0.007836 0.283230039 0.356109207 cg15665928 LINC01391 S_Shelf Body 7.62384 0.000159 0.365917357 0.467478253 cg15706250 ANK1 Island Body 94.2096 1.34E−10 0.403852326 0.520685646 cg15710198 SOX17 Island TSS200 6.19041 0.001184 0.205206583 0.260581234 cg15718581 ZFP42 N_Shore TSS1500 7.3251 2.19E−05 0.55978453 0.695412483 cg15724256 EN1 N_Shelf Body 4.36895 2.69E−05 0.251691757 0.388584155 cg15736169 LINC00403 Island TSS1500 8.55048 1.52E−06 0.123484738 0.233093171 cg15741433 HIST1HIA N_Shelf TSS200 4.2225 0.002514 0.429481923 0.53057402 cg15772924 HOXC6 S_Shore 5URT 4.56126 0.0031 0.550127569 0.629434141 cg15778437 PAX6 Island TSS200 6.13217 1.11E−05 0.062314925 0.16223126 cg15792487 CHRNB1 Island TSS200 6.78428 2.42E−05 0.225336556 0.334932777 cg15798385 EVX1 S_Shelf TSS1500 5.56661 0.000161 0.498128571 0.61769602 cg15826897 EPHX3 S_Shore 5URT 15.976 1.23E−11 0.320768647 0.539226052 cg15829969 NBPFS Body 4.25776 0.007346 0.641966651 0.692504707 cg15835620 ALDOC 5URT 51.9507 1.34E−09 0.439898614 0.605879393 cg15854847 NKX2-6 Island 1stExon 16.8258 9.97E−07 0.106743251 0.199136975 cg15861585 DRD4 Island TSS1500 21.3068 1.13E−09 0.28276645 0.451054231 cg15873301 SYN2 N_Shore TSS1500 4.54276 0.00062 0.20828984 0.324050357 cg15877520 RNF39 Island Body 5.33165 4.87E−06 0.252887085 0.416379843 cg15880760 CHAD Island TSS200 5.20142 0.000481 0.301170185 0.421657018 cg15911153 PROB1 Island 1stExon 20.8658 4.56E−06 0.602955 0.730096434 cg15912800 HOXA10-AS Island Body 4.31912 0.000214 0.275885303 0.398816031 cg15927196 PON3 Island TSS1500 24.4511 2.54E−08 0.397064698 0.513895455 cg15949044 PCDHGC4 Island TSS200 4.16741 2.39E−05 0.285209646 0.452341853 cg15951188 KCNAB8 N_Shelf 1stExon 12.4082 0.000496 0.725545721 0.793964232 cg15959715 GAD2 Island Body 7.60668 2.64E−06 0.128268286 0.221494658 cg15981851 AGAP1 Island Body 19.4335 1.49E−05 0.484664752 0.573708192 cg15982700 HOXA6 S_Shore 5URT 4.62446 0.003081 0.342638162 0.392987474 cg15985184 IGLON5 Island Body 14.7563 1.44E−07 0.438496219 0.596028991 cg15985775 DBX1 N_Shore TSS1500 4.44725 0.000598 0.32861342 0.430467044 cg15990629 BCAT1 N_Shore Body 11.5349 0.001263 0.69204914 0.738727727 cg16017144 RBFOX3 Island Body 12.6246 4.79E−05 0.557372422 0.677758396 cg16021909 TRIM58 Island Body 24.1732 1.29E−10 0.091625971 0.19752998 cg16030177 IRF6 N_Shore 5URT 22.1456 0.000129 0.65754907 0.733656909 cg16041686 PITX2 N_Shore 5URT 7.11277 8.23E−06 0.150582173 0.266662949 cg16043357 VAX1 N_Shore TSS200 9.71721 1.54E−07 0.216716208 0.351398275 cg16051228 TERT Island Body 7.07452 0.000551 0.363092129 0.432070229 cg16078649 RNF39 Island Body 12.6216 1.84E−06 0.542516702 0.662611394 cg16084872 TMPRS52 Island 5URT 4.03062 0.002127 0.649815595 0.462388268 cg16092786 WT1 N_Shelf Body 4.88907 0.000246 0.194023822 0.292325797 cg16104915 HOXA9 Island TSS200 7.23797 1.50E−06 0.110442312 0.224542336 cg16113681 TBX1 Island Body 6.87756 0.000196 0.26590667 0.364432953 cg16117799 PRDMS N_Shelf 5URT 4.93767 0.000679 0.216674095 0.302150582 cg16138458 HIST1H3I Island TSS1500 24.3189 1.74E−11 0.235883152 0.401079916 cg16180353 PAX6 N_Shore Body 24.1691 5.11E−09 0.478553853 0.63154397 cg16184495 EPHX3 S_Shore 5URT 20.8256 1.50E−11 0.374622083 0.595299737 cg16197925 FEZF1 Island 1stExon 7.93342 1.26E−07 0.175488396 0.305893535 cg16243019 DLX6 S_Shore Body 8.47873 1.65E−05 0.379708156 0.479205829 cg16246169 SOX2-OT N_Shore Body 11.1404 0.00043 0.651898698 0.735388485 cg16246489 H2AFY S_Shore TSS1500 5.22965 0.00075 0.522076207 0.620088596 cg16264705 ATP5G2 Island TSS1500 3.64373 0.00089 0.130249609 0.2355965 cg16281276 CYB5R2 Island TSS1500 6.16364 1.07E−06 0.195221381 0.339427944 cg16297569 GFI1 Island TSS1500 12.4415 7.13E−07 0.394518856 0.530127524 cg16302441 POMC Island TSS1500 16.1045 2.52E−06 0.58060109 0.712357444 cg16306115 KAAG1 Island 1stExon 10.0876 4.04E−08 0.136958562 0.263514457 cg16306978 APOB Island TSS200 3.8232 0.001406 0.51776577 0.632776777 cg16331823 FLOT1 Island TSS1500 10.0525 1.01E−09 0.267179987 0.470606525 cg16353006 LHX9 S_Shore Body 7.45714 1.50E−06 0.146044313 0.256929939 cg16362592 TBX20 N_Shelf ExonEnd 3.9498 0.005237 0.248790402 0.32555874 cg16368146 HCG9 N_Shore Body 19.533 2.03E−06 0.382086505 0.533255177 cg16373229 FEZF1 N_Shore Body 15.2909 4.74E−06 0.24131424 0.325261333 cg16374999 PRPH Island 5URT 4.34004 0.004208 0.642138028 0.738408418 cg16379704 ICA1 Island TSS1500 23.5199 2.37E−07 0.487858078 0.637063657 cg16390060 NKX2-2 Island 1stExon 11.3066 1.17E−07 0.160368595 0.265710746 cg16392213 FEZF1 Island Body 10.7743 3.20E−08 0.12859394 0.257871147 cg16407471 MIR129-2 Island TSS200 20.3473 6.36E−13 0.155951964 0.30999726 cg16410706 BATF3 Island TSS1500 6.92078 7.30E−08 0.14083266 0.321316707 cg16419441 DPYS Island Body 14.7332 8.32E−11 0.285184412 0.463370934 cg16435779 GRHL2 N_Shore TSS1500 6.43369 4.13E−06 0.197280159 0.331604184 cg16440561 SPEG Island Body 4.86042 0.000301 0.601825258 0.731216093 cg16458438 TBX5-AS1 Island Body 8.91599 2.61E−06 0.137558991 0.219814876 cg16459364 NEFM Island TSS1500 16.3338 3.48E−12 0.148411597 0.3423751 cg16463460 WT1 S_Shelf Body 8.13919 0.002365 0.599164116 0.660481455 cg16473141 REC8 Island 1stExon 29.4907 3.98E−11 0.402290596 0.613716657 cg16493531 FLOT1 Island TSS1500 7.65465 6.22E−08 0.37106665 0.591498945 cg16501028 WT1 S_Shore Body 10.6347 0.000218 0.412398791 0.482729667 cg16513459 KCNABB N_Shelf TSS200 11.2953 7.75E−06 0.643410566 0.764900253 cg16524928 NKX2-2 Island TSS200 24.504 1.95E−08 0.22039776 0.327503137 cg16529477 PAX3 N_Shelf TSS1500 8.77119 4.75E−07 0.444271716 0.583110137 cg16544169 NKX2-6 Island TSS1500 26.0301 5.67E−08 0.356898705 0.468082323 cg16546442 PITX2 Island Body 34.572 2.47E−12 0.227935612 0.354180323 cg16555165 SHF S_Shore 5URT 14.2226 2.28E−09 0.098624138 0.238725518 cg16557173 HOXB13 Island Body 8.46004 5.93E−07 0.253898867 0.393296651 cg16568727 TTBK1 Island Body 5.02281 0.008662 0.38536414 0.471324777 cg16574871 GDF6 N_Shelf Body 18.8817 6.91E−07 0.551820059 0.668923232 cg16582419 RNF39 N_Shelf Body 23.8595 2.91E−07 0.577653147 0.692054202 cg16604516 FBLN2 Island TSS1500 5.64401 0.00037 0.25504052 0.338056863 cg16615829 HSPB6 Island 3UTR 8.78791 3.92E−05 0.402149002 0.495262939 cg16615954 PAX6 Island Body 16.8774 2.24E−09 0.246096425 0.395636425 cg16620525 LVRN Island TSS200 18.9663 3.52E−09 0.116536159 0.228274784 cg16638920 LINC00461 Island Body 9.95991 2.58E−06 0.345610054 0.449626253 cg16642284 FOXI2 Island TSS200 4.94409 1.66E−06 0.159498339 0.335120386 cg16651126 HOXA4 Island TSS200 12.7467 1.77E−06 0.279915538 0.369349424 cg16652651 AMPD3 Island TSS200 4.27463 0.003605 0.119407053 0.186505427 cg16670554 SKOR1 Island Body 6.2874 6.17E−05 0.187367455 0.272957847 cg16686158 HORMAD2 Island TSS1500 24.628 3.44E−10 0.426714084 0.537359606 cg16688483 GRHL2 Island Body 9.96497 3.51E−06 0.328346919 0.455555121 cg16699148 TACSTD2 Island TSS200 10.1891 6.32E−07 0.404067054 0.532669444 cg16705627 LINC00403 Island Body 15.4489 4.37E−10 0.288392853 0.465216825 cg16723488 APOB Island TSS200 5.45366 0.00029 0.394224983 0.506687579 cg16742875 NKX3-2 N_Shelf Body 21.3459 0.0001 0.590751543 0.668214303 cg16747564 SOX1 S_Shelf TSS1500 6.46694 0.000228 0.180541206 0.255598232 cg16748008 HOXA3 Island 5URT 6.83301 0.000649 0.329235195 0.399208375 cg16758800 DLX6-AS1 S_Shore Body 20.0438 4.72E−10 0.31897493 0.43515604 cg16763443 TTC6 Island Body 18.7099 1.27E−08 0.384153504 0.501827192 cg16783744 DPYS Island Body 26.7207 1.00E−10 0.111274247 0.2171099 cg16786703 ADAM8 Island Body 8.11996 1.26E−05 0.229733819 0.335198744 cg16737431 HOXB9 S_Shore Body 5.75506 9.49E−05 0.457385319 0.569377566 cg16793594 CTX2 S_Shore TSS1500 5.84726 3.55E−05 0.205871502 0.307350709 cg16809460 DMRTA2 Island Body 3.84469 0.001442 0.16366241 0.23535763 cg16850930 SOX17 Island Body 4.80678 0.008074 0.434181574 0.500956586 cg16851417 SIM2 N_Shelf Body 10.1622 1.80E−07 0.202218953 0.323891397 cg16865446 PAX6 N_Shelf Body 9.78659 1.43E−10 0.360377724 0.587807573 cg16865298 WNT16 N_Shore Body 14.9448 9.24E−09 0.345177309 0.503736758 cg16891104 UPK3A Island Body 37.4026 9.71E−07 0.492674093 0.590980475 cg16957769 HOXC4 N_Shelf TSS1500 3.1423 0.004554 0.225776692 0.324842721 cg16933805 DBX1 Island TSS200 3.36172 0.009801 0.263865972 0.523280127 cg16961816 DLX6-AS1 N_Shore Body 4.69147 0.003976 0.398740023 0.470510556 cg16964025 PTPRN2 Island Body 8.76891 7.89E−09 0.198294818 0.380152949 cg16964348 NPY Island TSS200 11.7172 2.54E−10 0.115642753 0.252968067 cg16991515 HIST1H23K N_Shore 3UTR 4.248 0.001901 0.371044831 0.473967295 cg17011709 CYP26C1 Island Body 7.17817 1.64E−05 0.250935577 0.368466905 cg17019053 M1AP Island TSS200 3.83161 0.001744 0.267440985 0.34726932 cg17029062 TRIM71 Island TSS200 14.4008 1.39E−07 0.116275753 0.214411303 cg17029168 NKX2-2 Island 1stExon 8.1321 0.000164 0.127972987 0.19253687 cg17067528 IER3 S_Shore TSS200 7.5979 1.11E−05 0.29573945 0.444713377 cg17083494 DLX5 S_Shelf TSS1500 3.57615 0.008152 0.524175119 0.604749344 cg17107156 CHRN51 S_Shore Exon3nd 6.4386 0.000429 0.393853808 0.506406816 cg17158769 VAX1 Island Body 7.47368 2.29E−08 0.14756008 0.286338362 cg17153568 SP9 N_Shore TSS1500 5.27972 0.001353 0.143906081 0.204401204 cg17163168 TP73 Body 5.47612 0.000436 0.195054477 0.285020183 cg17181022 CYP26O1 S_Shore Body 8.10163 0.000254 0.328567705 0.410208475 cg17194154 LOC101927248 S_Shelf Body 8.0749 5.04E−07 0.83094332 0.191760647 cg17204275 SATB2 Island 5URT 4.30077 0.002371 0.17408703 0.254568259 cg17210938 TACSTD2 Island TSS200 5.46161 0.001427 0.219481331 0.283830022 cg17214381 ZFP42 Island TSS1500 21.8832 9.43E−14 0.25236621 0.447477493 cg17236169 NRXN2 Island Body 10.1161 1.91E−08 0.142049216 0.28691151 cg17241510 BARHL2 Island TSS200 12.9545 1.28E−08 0.364281279 0.518114808 cg17242957 PITX2 S_Shore 5URT 3.06255 0.008912 0.292894278 0.346999056 cg17280740 PAX6 S_Shore TSS1500 23.5664 1.60E−10 0.319711792 0.464824888 cg17298275 ABR S_Shore Body 4.53081 0.000533 0.590716659 0.697514659 cg17301223 CPLAH Island Body 3.75423 8.37E−05 0.273986153 0.423256957 cg17303540 PCDHGA8 S_Shore Body 3.91222 0.004776 0.591401333 0.661568626 cg17338212 LINC01891 N_Shore TSS200 26.7677 2.58E−08 0.312696 0.434859636 cg17353412 HOXA2 N_Shore 1stExon 6.05398 3.95E−05 0.468990587 0.59963218 cg17360661 SIM1 N_Shore TSS1500 7.78889 1.27E−06 0.131196166 0.241811092 cg17386213 NR2E1 Island TSS1500 5.62454 2.46E−05 0.396139808 0.517192194 cg17423207 JAK3 S_Shore TSS200 7.48503 0.000997 0.303975091 0.388413281 cg17452615 KDM2B S_Shore TSS1500 3.0564 3.16E−05 0.169394278 0.276190045 cg17462200 TBX5 N_Shelf 5URT 4.19652 0.001069 0.164499394 0.253568195 cg17476026 EPHX3 S_Shore 5URT 24.1894 5.18E−13 0.230741959 0.440025605 cg17486860 WT1 S_Shelf Body 7.83563 1.70E−06 0.322646384 0.474772945 cg17487170 TCF7 Island TSS200 18.4589 1.05E−06 0.523662933 0.62002597 cg17507671 PITX2 Island Body 10.5903 2.03E−06 0.221260498 0.323759115 cg17526485 EVX1 N_Shelf TSS1500 14.0883 1.08E−06 0.38908107 0.507617828 cg17561417 FBLN2 Island TSS200 6.21456 0.000107 0.195414183 0.284386715 cg17576288 POU4F1 N_Shelf TSS200 10.3282 7.05E−06 0.134618242 0.233541237 cg17594860 AKR7A3 Island TSS200 7.21862 1.80E−07 0.288662004 0.501674064 cg17602451 BCL2 Island Body 4.98574 0.000692 0.264673189 0.387334022 cg17611674 GATA3-AS1 Island Body 8.37078 3.09E−05 0.412713357 0.541990597 cg17626405 MAL Island Body 19.9531 1.39E−11 0.15047924 0.272047534 cg17630392 NPR3 Island 1stExon 21.7926 1.62E−13 0.257257505 0.481289699 cg17632937 HORMAD2 Island TSS200 18.9925 3.86E−08 0.305976708 0.445208051 cg17652435 HNF1B Island Body 8.5397 1.27E−05 0.112947249 0.211580814 cg17658976 PTPRN2 S_Shore Body 4.47817 0.004582 0.474659015 0.558749929 cg17665193 GAD2 Island Body 4.07202 0.002739 0.196090298 0.267172685 cg17674725 LYPD1 N_Shore 5URT 16.7602 5.96E−08 0.381969788 0.524156516 cg17694792 TCF24 Island 5URT 19.5938 7.67E−09 0.298240317 0.442733149 cg17696160 PAX9 Island Body 7.24553 6.94E−05 0.10366358 0.189356726 cg17698295 OPLAH Island Body 4.00191 2.86E−05 0.295852409 0.464395024 cg17727529 C1orf61 TSS1500 6.75367 0.00475 0.253236421 0.331276952 cg17737681 DLX1 S_Shelf Body 11.4466 1.97E−08 0.252875965 0.400162921 cg17753347 LHX9 N_Shelf Body 6.79502 1.54E−05 0.362180884 0.48494603 cg17768491 SPTBN4 Island Body 31.9516 2.57E−06 0.596831866 0.75167896 cg17774559 IRX4 Island Body 7.63032 1.52E−07 0.147490686 0.26915986 cg17799053 PTPRN2 N_Shore Body 16.5949 5.30E−05 0.598189566 0.680222495 cg17808882 FGF20 S_Shore TSS1500 11.3507 0.000247 0.517911349 0.592594455 cg17820365 GDF6 Island Body 4.9372 9.34E−05 0.209944898 0.32733554 cg17833476 TLX3 Island TSS200 21.7231 1.78E−09 0.174424072 0.281603325 cg17858325 GFI1 Island TSS1500 7.47149 1.14E−06 0.337729562 0.50938223 cg17883033 LYNX1 Island 5URT 11.9498 1.32E−05 0.58728876 0.488255848 cg17890928 CHAD Island 5URT 12.0383 4.17E−08 0.187541424 0.517844423 cg17894318 SOX2-OT N_Shelf Body 4.85505 0.006927 0.292918 0.351363888 cg17911021 SHF S_Shelf TSS200 47.9817 1.16E−09 0.466465321 0.645995889 cg17953764 ZAR1 Island 1stExon 12.1779 4.46E−08 0.350699606 0.489734795 cg17955729 PRRT1 Island Body 9.11046 0.002102 0.590286395 0.659701818 cg17968795 KIAA12110 Island TSS1500 12.1468 5.46E−10 0.340340157 0.546382612 cg17985645 TBX20 Island TSS200 19.8363 1.06E−14 0.105382315 0.285893297 cg17999576 GDF6 N_Shore Body 7.17236 6.61E−07 0.080711496 0.194295983 cg18002447 TBX2 N_Shelf Body 9.52199 2.55E−05 0.502601525 0.626567455 cg18004701 PTPRN2 Island Body 60.3432 1.31E−05 0.314220947 0.456252525 cg18008037 PTPRN2 Body 12.423 6.01E−05 0.515893961 0.601632879 cg18009690 EN1 N_Shelf Body 3.62404 0.006173 0.408456667 0.488124994 cg18020955 NBPF6 S_Shore Body 13.2146 9.68E−10 0.315350464 0.512824132 cg18034737 ZFP42 Island TSS200 15.1511 1.32E−11 0.298751612 0.528559364 cg18055747 BHMT Body 6.54294 6.07E−05 0.310740581 0.419380118 cg18065017 LHX4 Island Body 8.54376 2.43E−07 0.164405681 0.288955422 cg18064631 PDX1 N_Shore Body 15.0095 1.43E−08 0.305634029 0.441545632 cg18082638 PAX6 Island Body 12.9733 6.76E−11 0.232736846 0.404154903 cg18087266 RUNX3 Island TSS1500 8.07854 0.000874 0.427078248 0.498311343 cg18097224 TCF7 Island TSS200 11.7289 3.81E−09 0.297002078 0.489776421 cg18106668 NPY N_Shore TSS1500 6.60369 2.63E−05 0.208961779 0.315276483 cg18128164 JSRP1 Island Body 6.93476 5.35E−06 0.242165326 0.370465384 cg18144593 LINC01145 Island Body 5.24299 0.001173 0.291878907 0.365592041 cg18181229 PBX1 Island Body 19.1988 1.66E−05 0.709799167 0.815114051 cg18182148 GFI1 Island TSS1500 24.0419 6.65E−06 0.439434638 0.532292707 cg18182392 DES N_Shore 5URT 6.69962 1.42E−06 0.182051221 0.331384302 cg18202448 ELAVL4 Island TSS200 10.8587 5.35E−08 0.078857392 0.173124169 cg18221467 GFI1 Island 5URT 3.32963 0.00269 0.098510456 0.181995669 cg18229178 HSPB5 Island 3UTR 6.31466 5.39E−05 0.491426869 0.627269212 cg18235100 PRDM8 Island Body 5.33123 9.01E−06 0.225875627 0.358635163 cg18240703 RBFOX3 S_Shore Body 8.59428 0.000115 0.290458174 0.387115047 cg18247055 SPAG6 Island TSS200 19.3193 1.02E−14 0.124712281 0.338985005 cg18267374 NEFM Island TSS1500 13.4286 1.04E−08 0.20676212 0.333002336 cg18304448 LOC90768 N_Shore Body 26.9983 0.000597 0.714294194 0.773667212 cg18311537 HOXA10-AS Island Body 5.38304 0.002059 0.254178464 0.327363658 cg18318878 PROX1-AS1 Island Body 13.299 9.34E−08 0.187029062 0.298427348 cg18322569 BARHL2 Island 1stExon 6.29987 2.16E−08 0.197396521 0.353766007 cg18327157 PTPRN2 Island Body 13.4436 4.96E−05 0.61934575 0.708686101 cg18336674 TLX3 Island TSS1500 9.73175 8.58E−07 0.298246398 0.407585068 cg18342279 ZAR1 Island 1stExon 30.9515 5.65E−12 0.062975676 0.17502701 cg18358723 LINC00461 N_Shore Body 26.2231 2.01E−13 0.360938969 0.556964268 cg18362330 NEUROG1 S_Shore TSS1500 4.73319 0.002724 0.352742707 0.415974497 cg18366919 EPHX3 Island TSS1500 59.53 2.32E−14 0.401490765 0.628729481 cg18372607 PAX6 S_Shore TSS1500 12.5926 1.60E−08 0.419495719 0.620407781 cg18394840 H2AFY S_Shore TSS1500 9.82185 7.07E−07 0.22672006 0.327011825 cg18400845 DKFZp686K1684 Island TSS200 7.81442 0.000245 0.160146836 0.239914486 cg18405727 TRIM71 Island 1stExon 10.2963 3.19E−09 0.073522285 0.195150408 cg18406197 FBLN2 Island TSS200 5.70911 0.000326 0.115995684 0.196359008 cg18407955 PTPRN2 Island Body 4.19021 0.001203 0.250637437 0.349141454 cg18416576 HOXA10 Island TSS200 13.6841 9.58E−08 0.121160838 0.21969374 cg18435852 GATM S_Shore TSS200 4.08219 0.002803 0.472096821 0.568737838 cg18447772 HOXA9 Island TSS200 6.5081 9.51E−07 0.167529767 0.285976453 cg18459489 VAX1 N_Shore 3UTR 7.67101 5.78E−05 0.248709798 0.352316234 cg18495682 RGCC Island Body 10.0128 3.08E−06 0.253291282 0.38447752 cg18501555 PITX2 Island 5URT 7.90439 4.41E−07 0.164471828 0.268454837 cg18502142 DLX6-AS1 Body 3.81352 5.22E−09 0.356040295 0.528492931 cg18507579 FCDHGB6 N_Shore TSS1500 10.4458 9.58E−10 0.052228509 0.233778193 cg18512945 RECS Island 5URT 32.5159 2.72E−09 0.493475405 0.690941253 cg18528367 SIM1 Island TSS1500 5.82481 0.00082 0.151508981 0.21755899 cg18534491 RECS Island TSS200 24.0735 5.41E−12 0.305632671 0.502323024 cg18539122 RNF219-AS1 N_Shore Body 19.1792 1.57E−10 0.266935236 0.335809937 cg18544365 MIR199A1 TSS200 28.236 2.54E−05 0.708369473 0.825552071 cg18552361 GDF7 Island TSS1500 7.40243 2.88E−09 0.232560626 0.42899634 cg18556834 REC8 Island TSS200 30.6884 3.98E−12 0.317846961 0.518107681 cg18560204 BNC1 Island TSS1500 16.7946 2.51E−05 0.396627543 0.478962843 cg18569734 NR281 N_Shore Body 16.3179 5.93E−10 0.239844657 0.381504319 cg18579879 WNT16 N_Shore Body 19.4183 5.84E−10 0.289589062 0.425330533 cg18588323 TNFAIP8L3 Island Body 5.8607 6.37E−06 0.457123502 0.627573555 cg18599069 GATA3 Island 5URT 5.18647 0.00034 0.318744713 0.418446436 cg18601426 PTPRN2 Island TSS1500 7.4007 0.000319 0.378384829 0.470086596 cg18603228 FBLN2 Island TSS200 4.1224 0.000785 0.175258153 0.269520406 cg18609783 SP9 N_Shore TSS1500 5.93052 9.08E−05 0.199823974 0.292209257 cg18614735 ANK1 Island Body 5.56832 0.000796 0.549223589 0.636190121 cg18617005 PCDHGB6 N_Shore TSS1500 8.11405 2.83E−09 0.115357692 0.28455606 cg18628371 REC8 Island TSS200 26.4715 4.77E−14 0.247613071 0.447591304 cg18650716 FLOT1 Island TSS1500 13.1511 6.64E−11 0.167437264 0.36048667 cg18664869 PCDHB15 S_Shelf 5URT 5.50884 3.78E−05 0.162265126 0.27620882 cg18687675 GSC Island Body 15.3921 7.49E−10 0.291807365 0.470013512 cg18691434 GPC2 Island TSS1500 15.5553 3.08E−07 0.257974009 0.358204774 cg18725076 CTNND2 S_Shore TSS1500 23.1262 1.14E−08 0.118424491 0.193368514 cg18736063 NKX2-4 S_Shore TSS1500 5.0295 0.000205 0.182061271 0.282565111 cg18760597 FAM184B Island TSS200 9.11841 1.22E−09 0.070912743 0.237619076 cg18795809 ZNF518B Island 5URT 8.36665 5.57E−07 0.129595157 0.240899469 cg18796287 PROB1 Island TSS200 20.5923 1.29E−08 0.539499651 0.701972818 cg18817990 NKX3-2 N_Shore 3UTR 5.31089 0.003429 0.568214085 0.636472949 cg18867200 LINC01145 S_Shore Body 4.77564 0.006418 0.520552419 0.594372525 cg18867923 TBX1 S_Shore Body 11.9792 6.97E−06 0.427310295 0.546285681 cg18873386 DLX5 S_Shore Body 7.51178 4.30E−05 0.490907524 0.599955256 cg18884137 CHRNB1 Island 1stExon 7.60404 1.06E−06 0.183831358 0.316830618 cg18898125 NEFM N_Shore TSS1500 9.61316 2.33E−06 0.437298423 0.598517638 cg18943485 LBX1-AS1 S_Shore Body 16.5964 8.12E−10 0.37951718 0.591820949 cg18950773 WT1 S_Shelf Body 15.1075 8.25E−09 0.237997974 0.364574508 cg18952647 BNC1 Island TSS1500 5.37501 9.25E−05 0.376393719 0.498551567 cg18984724 SPEG Island TSS200 4.16953 0.008523 0.421400391 0.489853415 cg19006378 PAX6 S_Shelf 5URT 14.5559 4.86E−06 0.191494257 0.287171432 cg19006429 RNF69 Island 1stExon 5.6169 9.01E−06 0.262171297 0.411383047 cg19022697 TTC22 Island Body 12.4534 9.76E−06 0.376542029 0.461752303 cg19043574 KIAA1211L Island Body 5.73261 1.35E−06 0.226254804 0.402278031 cg19046825 CHAD Island TSS200 4.13726 0.000578 0.27575857 0.410218347 cg19092981 TBX1 Island Body 12.4816 0.000116 0.720006729 0.813349848 cg19098763 ELAVL4 Island TSS200 10.6067 7.11E−10 0.084956398 0.221883706 cg19103219 LINC01475 N_Shore Body 4.12703 0.008564 0.26200805 0.310319771 cg19113641 GUF7 N_Shelf TSS200 4.52751 0.000657 0.055786848 0.132334285 cg19127283 TRIM71 Island TSS200 11.8564 1.03E−10 0.126788553 0.281658589 cg19181476 NRXN2 Body 11.2077 1.69E−05 0.437986124 0.543260569 cg19164987 HOXC6 Island 5URT 3.45532 0.00477 0.512598324 0.591719046 cg19181162 TTC6 Island Body 21.4468 1.09E−10 0.294844788 0.448929216 cg19183743 HOXA-AS3 S_Shore Body 5.58012 0.000165 0.382559529 0.493138218 cg19211915 WT1 Island TSS200 21.0418 2.34E−10 0.247232195 0.391269267 cg19241327 PTPRN2 Island Body 8.50737 2.28E−06 0.329177162 0.452448658 cg19247475 PCDHG36 Island 1stExon 44.3066 8.64E−09 0.611487473 0.770575283 cg19255034 SPTBN4 Island Body 4.99719 0.003627 0.522056947 0.594846242 cg19270505 RUNX3 Island 1stExon 7.77555 8.12E−07 0.113755246 0.218348874 cg19281363 RGCC Island TSS1500 16.0748 2.15E−06 0.352716495 0.503901001 cg19290410 TBX5-AS1 Island Body 7.32905 1.35E−06 0.147292054 0.253437205 cg19305681 CYP26C1 N_Shelf Body 4.90528 0.000157 0.153316146 0.261350188 cg19325985 HOXD18 S_Shore TSS1500 9.1827 1.87E−06 0.399982423 0.534120051 cg19329160 RNF219-AS1 Island Body 8.60088 1.98E−06 0.164899784 0.267264503 cg19334350 PTPRN2 S_Shore Body 9.22478 4.00E−06 0.314449187 0.444292523 cg19363499 PRDM14 Island 5URT 11.6897 5.45E−06 0.142179275 0.226016956 cg19378036 HNF15 Island TSS1500 4.34373 7.99E−05 0.266627511 0.592619205 cg19384289 HOXD8 Island TSS200 6.01803 1.19E−05 0.099368389 0.2095155 cg19392656 F5LN2 Island TSS1500 5.1189 9.12E−05 0.15974956 0.248074327 cg19406029 SOX7 S_Shore Body 10.382 3.83E−07 0.453441531 0.625600202 cg19403104 RCN8 Island 1stExon 6.43763 1.04E−05 0.10711464 0.214153279 cg19407095 SOX1 Island TSS200 15.6357 0.000156 0.215413566 0.282166094 cg19410791 CD8A S_Shelf TSS1500 6.76389 0.003426 0.659376005 0.735806919 cg19416570 ZNF247 Island 5URT 18.2918 2.64E−08 0.311616852 0.411936929 cg19497031 POU4F1 Island Body 20.3991 9.86E−13 0.08002373 0.204831693 cg19498960 SIM2 N_Shelf Body 6.4801 0.000118 0.213305814 0.306683987 cg19520234 MGAT4D Body 6.33852 0.000227 0.426039488 0.500163212 cg19537719 ANK1 Island Body 7.12919 0.00015 0.622529761 0.752024889 cg19544662 FOXI3 Island TSS200 11.033 3.26E−10 0.221597176 0.599217105 cg19578835 TTC6 Island 5URT 8.40703 7.02E−09 0.125352756 0.267933818 cg19593490 HCG9 N_Shore Body 7.79059 0.000178 0.369764078 0.480599628 cg19594666 LEP Island TSS200 7.6217 3.16E−05 0.287823723 0.591379455 cg19616230 SLC34A2 Island TSS1500 6.6873 7.48E−05 0.301621431 0.40064544 cg19617672 WNT16 N_Shore Body 8.89686 1.27E−05 0.132662698 0.22709548 cg19628148 CCDC37 Island 5URT 4.79564 0.000186 0.159824364 0.251712756 cg19689322 KCNQ4 S_Shore Body 8.22529 6.89E−06 0.360362426 0.437983049 cg19711783 NTN1 Island Body 12.3025 5.86E−14 0.15382545 0.417709614 cg19718882 WT1-AS N_Shore Body 5.72747 4.37E−05 0.212520009 0.325131857 cg19725343 IGLON5 Island Body 5.59984 1.08E−06 0.289951525 0.475914244 cg19733042 NBPF8 Body 6.81366 0.005675 0.701039426 0.761447444 cg19741645 EBF3 N_Shore Body 16.5096 0.000135 0.736955496 0.797555616 cg19741945 TRIM71 Island 1stExon 16.574 2.45E−10 0.071768964 0.719200876 cg19759671 LOC90768 Island Body 7.87859 0.000317 0.594065572 0.714661078 cg19760241 LHX1 Island Body 12.3234 2.83E−08 0.215127779 0.338894594 cg19761110 AGAP1 Island Body 12.8399 2.15E−06 0.459220209 0.582516 cg19762657 MAL S_Shore Body 5.40415 0.00039 0.188733259 0.274674552 cg19766441 SLC34A2 Island TSS1500 7.50814 2.64E−07 0.176200335 0.330598912 cg19767800 NR2F1-AS1 N_Shore Body 26.8457 2.91E−08 0.547704163 0.74524497 cg19770541 ZAR1 N_Shore TSS1500 8.06612 0.000348 0.318610147 0.381747705 cg19792544 KIFC2 N_Shelf 5UTR 15.4103 7.71E−07 0.298568269 0.448074728 cg19802133 LINC00403 Island Body 29.1631 9.82E−11 0.344062543 0.496177364 cg19808973 RCN3 Island 1stExon 6.64664 9.10E−07 0.087848633 0.221149819 cg19816811 HOXA-AS3 N_Shore Body 3.84325 0.002921 0.49767724 0.589245283 cg19842809 PAX9 S_Shore TSS1500 5.90394 0.000151 0.236785531 0.324544787 cg19844325 ANK1 S_Shore TSS1500 10.6964 1.36E−06 0.208115856 0.519461275 cg19844653 TCF7 S_Shore TSS200 11.7742 0.000104 0.420360022 0.504726748 cg19852958 NKX3-2 Island 1stExon 11.0387 3.63E−10 0.213761278 0.390592106 cg19876672 VAX1 N_Shore 5UTR 5.92333 1.02E−05 0.267581817 0.406519767 cg19884262 FOXI2 Island 1stExon 12.2074 4.67E−08 0.253363243 0.405949632 cg19909239 PITX2 Island 5URT 14.0877 2.05E−04 0.190493558 0.231078255 cg19923650 NKX2-5 N_Shelf 5UTR 7.63611 1.63E−07 0.120904266 0.256820906 cg19961043 CHAD Island TSS1500 5.85916 8.87E−05 0.198159036 0.311028029 cg19962750 DLX5 S_Shelf TSS1500 22.7974 2.38E−05 0.636280651 0.722495444 cg19980151 NR2F1-AS1 N_Shore Body 9.97179 3.06E−08 0.351144177 0.577298493 cg19989295 REC5 Island TSS200 220.461 9.87E−14 0.285248684 0.530623958 cg20003638 SLC12A5 S_Shore 5UTR 100.175 4.98E−05 0.724658581 0.847950172 cg20014393 PAX6 Island TSS200 19.3007 8.68E−14 0.36145828 0.620588756 cg20014822 FGF19 N_Shore Body 4.61986 0.000787 0.211106193 0.296092176 cg20027296 TLX3 Island Exon3nd 15.8661 6.96E−07 0.420871829 0.542703212 cg20049415 NKX2-4 Island 1stExon 35.0598 3.29E−12 0.15292705 0.283244342 cg20065463 DE5 N_Shore 5URT 7.10058 1.30E−07 0.272935362 0.451054271 cg20073553 NKX3-2 Island Body 5.77127 0.004553 0.424616469 0.511879401 cg20080624 DLX5 Island Body 4.99317 0.000399 0.45875945 0.533856923 cg20094850 PRAC2 N_Shore TSS1500 8.91098 9.65E−07 0.200785164 0.317781466 cg20099830 TBX5 N_Shelf 5URT 10.081 6.28E−08 0.310201287 0.458847958 cg20120208 TLX1 S_Shore 3UTR 3.81635 0.002147 0.301642671 0.376603652 cg20125091 GFI1 S_Shore TSS1500 29.536 2.12E−06 0.542301519 0.643652556 cg20146541 TRIM58 Island 1stExon 21.9197 4.31E−17 0.110349952 0.363890803 cg20177385 SLC41A3 TSS200 10.8829 2.00E−06 0.350382508 0.44587501 cg20204986 WT1 N_Shore Body 6.5651 0.000519 0.403866388 0.474765061 cg20232102 FPP1R14A Island 1stExon 12.0711 1.60E−10 0.175226145 0.362609666 cg20248516 EN1 N_Shelf Body 5.52909 7.46E−05 0.292354778 0.391654679 cg20249327 RNF39 Island Body 7.57819 4.008-06 0.285282178 0.423858529 cg20250981 KCNQ1DN Island TSS1500 4.28637 0.001209 0.349317588 0.439709273 cg20262306 SHH S_Shore 5URT 13.0136 8.04E−07 0.509360164 0.655855596 cg20264762 ESRP2 N_Shelf 1stExon 22.8949 3.08E−08 0.449200194 0.583696576 cg20272979 ITPKA Island Body 10.8957 1.45E−06 0.546541036 0.736952009 cg20293942 DLEU1 Island Body 23.9208 2.79E−06 0.621626008 0.719852111 cg20311863 BARHL2 Island TSS1500 14.0965 1.83E−08 0.094841735 0.200530401 cg20312228 CCDC37 Island TSS200 12.8881 2.06E−07 0.141265823 0.24330336 cg20322977 CYP26C1 Island 1stExon 6.50117 0.003475 0.417134062 0.481221879 cg20342079 BCAT1 N_Shore TSS200 7.3502 1.03E−06 0.234865636 0.367089444 cg20359285 EN2 S_Shelf 1stExon 7.63444 0.000125 0.06592167 0.142306297 cg20371765 ICA2 S_Shore TSS1500 10.2615 9.41E−09 0.269209276 0.442337595 cg20377305 DLX5 Island Body 8.39082 2.67E−07 0.150303543 0.264619552 cg20381115 GATM S_Shore TSS200 4.21907 0.004171 0.385953412 0.476913108 cg20387815 POMC Island TSS200 6.15625 2.36E−05 0.328587554 0.475714212 cg20399462 LINC00403 Island Body 4.98615 0.004884 0.246342644 0.299335357 cg20399871 HOXA9 Island 1stExon 3.72556 0.007223 0.204600265 0.269153764 cg20403557 LVRN Island Body 4.40426 0.000455 0.228286075 0.329090354 cg20429172 TRIM58 Island TSS200 16.8021 1.21E−11 0.20154422 0.366979731 cg20454586 TTBK1 Island Body 5.32912 0.000785 0.197848039 0.277545371 cg20467957 DKFZp686K1684 N_Shore Body 73.2361 5.81E−11 0.480357829 0.581849596 cg20469799 KATNAL2 N_Shore TSS200 7.23421 1.08E−03 0.120875516 0.286978555 cg20512711 DKFZp686K1684 Island Body 15.5295 5.50E−14 0.175467838 0.379902061 cg20521696 SHH S_shore 5URT 19.7959 4.30E−06 0.598288736 0.710110727 cg20528093 DKFZp686K1684 Island TSS1500 4.33392 0.001201 0.479566101 0.581024374 cg20529344 DLELI1 S_Shelf Body 10.5902 6.25E−05 0.546257504 0.632352505 cg20540209 ISL2 Island Body 5.18125 0.000186 0.12172766 0.169163155 cg20541178 LYNX1 Island 5URT 7.57694 0.0002 0.195868433 0.275332524 cg20572816 TLX1 N_Shelf Body 3.475 0.00267 0.23055551 0.314436154 cg20574490 GRHL2 Island TSS200 18.0176 3.25E−11 0.299093352 0.48818099 cg20592655 C1orf51 N_Shore 5URT 6.1528 9.44E−05 0.261635712 0.347146778 cg20615879 TLX1 N_Shelf Body 5.41754 2.17E−05 0.241452736 0.359267755 cg20650802 IER3 Island 1stExib 5.49158 2.84E−07 0.309682419 0.520558419 cg20672981 HOTTIP N_Shelf Body 26.5811 2.13E−07 0.487395504 0.597795364 cg20682145 PBX1 Island Body 31.5081 1.51E−07 0.490240953 0.589135818 cg20688917 HOXA3 N_Shore TSS1500 7.1669 0.004168 0.729594217 0.808881212 cg20705985 FITX2 N_Shore 5URT 5.48117 0.002743 0.156584745 0.207863097 cg20710709 PAX9 Island Exon3nd 5.02087 0.000623 0.365077054 0.456764564 cg20725013 HOXA-A53 S_Shore Body 6.70603 0.000115 0.130368186 0.223778848 cg20751469 NR2E1 N_Shore Body 9.86331 5.60E−06 0.42968854 0.563242816 cg20739435 LOC648987 S_Shore TSS1500 3.80035 0.00209 0.21265756 0.308835924 cg20747380 HOXA2 N_Shelf 1stExon 17.7323 4.10E−06 0.496721643 0.594180293 cg20755721 SIM2 Island TSS1500 5.34343 0.00409 0.221784242 0.291734221 cg20785796 SATB2 Island 5URT 9.01048 1.70E−06 0.381194418 0.497477384 cg20800022 GDF6 Island Body 10.9567 2.06E−07 0.159999512 0.280002285 cg20801476 EVX1 N_Shore TSS1500 7.87763 2.65E−05 0.328250075 0.440461091 cg20817131 HOXA5 Island TSS1500 4.18629 0.002962 0.421847659 0.502763475 cg20817902 PRDM5 Island Body 6.34751 7.24E−06 0.097698513 0.204762599 cg20884887 HOXA10-A5 N_Shore TSS1500 4.75543 3.31E−06 0.233650447 0.374850679 cg20892260 NKX2-6 Island 1stExon 20.4729 4.78E−07 0.145169552 0.232744898 cg20927242 HLA-F Island Body 9.88346 4.65E−08 0.18051507 0.335948479 cg20931042 DRC4 Island TSS200 3.4648 2.41E−06 0.09255981 0.191767053 cg20941110 BNC1 Island TSS1500 3.96019 0.006217 0.502537245 0.575406453 cg20956788 ONECUT2 Island Body 7.64958 5.00E−05 0.162875956 0.248624632 cg20959460 LOC101929154 N_Shore TSS1500 16.1241 1.71E−11 0.314473133 0.492731758 cg20974609 HOXA5 N_Shore Body 4.33753 0.001555 0.428540455 0.504480212 cg20977170 ICA1 Island TSS1500 12.7054 1.57E−08 0.092850781 0.227330972 cg20985073 ALDH1L1 S_Shore TSS1500 7.76164 0.000203 0.413372124 0.514290566 cg20992114 FITX2 S_Shelf Body 8.59712 0.000777 0.339611031 0.414515455 cg20994254 HOXA11-AS S_Shelf Body 3.25154 0.005757 0.227420309 0.293247925 cg20994660 TBR1 S_Shore TSS1500 4.37658 0.000809 0.40604368 0.507094444 cg21010475 SHISA3 S_Shore Body 8.41646 2.86E−05 0.262301315 0.357312472 cg21021139 CYB5R2 Island 5URT 8.9749 1.34E−06 0.397531471 0.567621456 cg21016855 PAX6 N_Shore Body 12.1428 4.73E−12 0.224624653 0.455485739 cg21017569 KCNA33 N_Shelf TSS200 12.4852 0.001121 0.800880837 0.878251333 cg21052682 FGF29 S_Shore Body 7.86271 1.99E−06 0.073381482 0.164926489 cg21065282 GSC N_Shelf Body 8.0954 1.61E−05 0.120703974 0.207035663 cg21063716 ASCL2 Island TSS1500 12.9965 2.31E−06 0.29074841 0.394816487 cg21073927 GATA4 Island 5URT 10.4951 5.61E−07 0.227256102 0.34987529 cg21073950 MAST1 Island Body 4.26334 0.005436 0.718901295 0.795161434 cg21101465 PDX1 S_Shore TSS1500 5.35582 0.00317 0.339050261 0.434711273 cg21113740 ADGRL4 Island 1stExon 9.1566 1.91E−08 0.282772483 0.44115089 cg21117330 PCDHGA8 N_Shore 5URT 5.08556 0.00041 0.476529093 0.580492727 cg21124497 TRIM71 Island TSS200 12.8291 7.26E−09 0.186335319 0.248641648 cg21125569 HOXD13 S_Shelf TSS1500 8.68744 6.63E−06 0.459708043 0.604441173 cg21132352 CHRN51 S_Shore Body 11.2103 3.30E−05 0.564250713 0.690790717 cg21134232 HOXA3 S_Shore 5URT 4.19316 0.002935 0.490188502 0.552001929 cg21146503 DBX1 Island Body 9.4313 0.000103 0.227824308 0.322576416 cg21165027 NKX2-3 Island TSS1500 13.0488 2.30E−06 0.29774207 0.392608053 cg21172532 BCAT1 N_Shore 2stExon 8.12604 9.53E−06 0.345576027 0.478118571 cg21172377 HOXA10-HOXA9 Island Body 8.81032 4.60E−06 0.219481758 0.322923671 cg21174746 CTNND2 N_Shore Body 8.03936 2.87E−06 0.176852071 0.292922422 cg21195468 LHX2 S_Shore TSS1500 7.65383 3.16E−05 0.222284622 0.322005837 cg21100656 NKX2-4 Island TSS200 15.2788 2.42E−08 0.168404029 0.289018852 cg21200703 SLC34A2 S_Shore 5URT 8.28113 5.52E−05 0.461891595 0.567723828 cg21201099 TBX2 Island Body 4.36298 0.001954 0.247275088 0.329164387 cg21211357 CPM Island TSS1500 12.8348 6.19E−06 0.341610504 0.445220564 cg21242356 HOXB-AS3 N_Shore TSS200 8.27475 1.89E−05 0.334272388 0.44095593 cg21245652 MAL Island TSS1500 19.5844 2.22E−07 0.307562458 0.416126319 cg21249371 KDM28 S_Shore TSS1500 3.36862 0.002204 0.31509408 0.443699404 cg21266502 LRAT N_Shore 5URT 6.17457 2.45E−07 0.167118575 0.323547329 cg21269897 HIST1H2BK N_Shore 3UTR 5.45511 0.000419 0.279402227 0.379044661 cg21279601 DRD5 N_Shore TSS1500 6.97504 0.00063 0.534905785 0.634019293 cg21303021 THRB-AS1 Island Body 4.024 0.000615 0.308635694 0.414030698 cg21429745 LBX2 Island Body 3.97637 1.13E−06 0.096486193 0.201253287 cg21460081 HCXB4 S_Shore TSS1500 11.8038 2.44E−09 0.084045451 0.199574331 cg21463349 EN1 N_Shore Body 11.365 5.99E−07 0.78455924 0.163290233 cg21472506 OTX1 N_Shore Body 5.45161 5.21E−05 0.210010069 0.419778793 cg21174786 HOXB3 Island 5URT 5.25595 0.007164 0.424891899 0.480124343 cg21484226 TRIM71 Island TSS200 13.2 7.02E−11 0.128590807 0.281500941 cg21485895 SATB2 Island 5URT 4.42512 0.000713 0.132935433 0.218265706 cg21561970 ISM1 Island TSS1500 8.86787 2.83E−08 0.120169377 0.263760053 cg21595709 EPHX3 Island TSS1500 81.9004 2.02E−15 0.303808994 0.497211734 cg21601837 ALDH1L1 S_Shore TSS1500 5.3971 0.000174 0.375285658 0.295886224 cg21606115 PTPRN2 Island Body 3.8577 0.000691 0.14111287 0.234996 cg21614303 PITX1 S_Shore Body 8.03165 9.19E−05 0.425736203 0.543282747 cg21652591 DLX5 S_Shore Body 4.31227 0.008673 0.250049517 0.312964366 cg21684012 SIM1 Island TSS1500 17.9987 1.26E−09 0.083035309 0.18335952 cg21697851 SIM2 N_Shelf Body 9.87866 2.25E−09 0.17714261 0.344302642 cg21711132 GABRG3 Island TSS1500 8.00195 9.13E−06 0.323187644 0.488412327 cg21713473 LRAT N_Shore 5URT 5.75518 3.99E−05 0.218598381 0.327757801 cg21726372 PITX2 S_Shore Body 5.04614 2.13E−05 0.146965164 0.252853967 cg21753226 KCNO1DN Island TSS1500 5.29155 4.53E−05 0.360042953 0.503972436 cg21762523 NR2E1 Island Exon8nd 4.27447 0.002898 0.103195074 0.161828936 cg21769619 WT2 S_Shelf Body 12.9888 7.56E−05 0.46072276 0.560838152 cg21774338 DMRTA2 Island 3UTR 8.93362 9.90E−06 0.279551432 0.585666273 cg21784388 ESRRG Island TSS200 12.0597 3.92E−09 0.224802477 0.356644401 cg21791017 LTK Island Body 21.3453 7.70E−07 0.44291415 0.556013485 cg21793948 BCL2 S_Shelf TSS1500 5.82334 2.97E−05 0.17768038 0.285786712 cg21811143 EN1 Island 3UTR 2.95187 0.004889 0.198834755 0.294231757 cg21843594 HORMAD2 Island TSS200 13.3142 1.29E−09 0.314167553 0.483250241 cg21865150 HOX513 Island Body 4.96682 2.03E−05 0.313784264 0.441979663 cg21890667 HORMAD2 Island TSS1500 26.5912 4.98E−12 0.424362917 0.587528212 cg21922060 LINC01391 N_Shore TSS200 6.30522 3.31E−06 0.376253791 0.529332651 cg21915513 PCDHGA8 Island 1stExon 24.5935 4.37E−09 0.566020798 0.735252939 cg21982368 SHF S_Shore 5URT 37.997 7.77E−10 0.463776553 0.640857477 cg21945930 DLX6-AS1 N_Shelf Body 5.13758 0.000253 0.339683889 0.428517229 cg21949512 TBR1 Island Body 5.75984 1.41E−05 0.099245161 0.19930497 cg21951975 IRF6 S_Shore TSS1500 20.498 3.69E−08 0.299056932 0.431778556 cg21971285 DKFZp686K1684 S_Shelf Body 21.6561 2.00E−10 0.28271755 0.421670949 cg22010052 LVRN Island 1stExon 15.6049 3.14E−10 0.194089848 0.347209183 cg22058124 TNFAIP3L3 Body 3.62007 0.00944 0.395430767 0.454771545 cg11039909 TP73 Island Body 12.1417 0.001467 0.760562326 0.83305797 cg22044002 CYP25C1 Island Body 5.90765 3.77E−05 0.384815199 0.588603907 cg22056595 PTPRN2 Body 6.13129 0.007287 0.333684517 0.398391444 cg22084642 DNAH1 5URT 6.65081 5.29E−06 0.418105356 0.562459404 cg22097249 BEND4 Body 5.54496 0.000239 0.43385394 0.534596382 cg22105332 RNF39 Island 1stExon 5.08268 7.14E−07 0.241261662 0.425499092 cg22118524 ESRRG Island TSS1500 11.5249 2.64E−08 0.120001963 0.229305397 cg22120446 LINC01391 N_Shore Body 9.3174 0.000149 0.48362691 0.579314172 cg22125838 WNT10A Island Body 7.22989 0.000129 0.260322178 0.365792475 cg22132508 LINC01391 S_Shelf Body 4.12412 0.0032 0.259183763 0.332785868 cg22149137 HOXB13 Island Body 7.48542 1.25E−05 0.247579345 0.381321621 cg22204479 NKX2-4 Island TSS200 6.01779 5.77E−05 0.122060921 0.236212891 cg22241820 SKOR1 N_Shelf Body 11.6758 0.000752 0.660270558 0.735255717 cg22249789 OTX1 S_Shore Body 8.56542 1.78E−09 0.173777348 0.353262294 cg22265644 NKX2-2 Island TSS200 12.7748 8.71E−07 0.257804912 0.355724495 cg22268510 PRRT1 Island Body 4.55532 0.003525 0.573636291 0.655524051 cg22272282 HIST1H2BK S_Shore 3UTR 5.48686 0.001937 0.561603825 0.656952361 cg22283925 ZFP42 S_Shore 5URT 11.1438 1.37E−06 0.433798837 0.552435273 cg22287064 MYO15B Island TSS200 7.13041 4.13E−07 0.110727003 0.241164653 cg22289831 SIM2 N_Shelf Body 3.4192 0.008976 0.232055198 0.313108439 cg22303418 OTX1 S_Shore Body 26.3033 6.03E−05 0.600733333 0.701248919 cg22341104 GFI1 Island 5URT 3.88568 0.000395 0.215481322 0.300320506 cg22345692 NPR3 Island Body 12.1034 1.10E−07 0.09600835 0.198956758 cg22436335 LHX4 S_Shore Body 6.16602 9.09E−05 0.311893325 0.42341982 cg22442454 IRF5 Island 1stExon 6.34926 1.63E−05 0.099544043 0.22550171 cg22459630 SKOR1 Island Body 5.80506 0.000303 0.191352767 0.274424614 cg22463553 PCDHG36 N_Shore 1stExon 14.249 4.73E−06 0.606110295 0.720875717 cg22469274 HOXA-AS3 Island Body 17.2804 6.47E−10 0.057619553 0.172769822 cg22474464 NKX2-2 Island Body 8.64867 1.14E−10 0.282635274 0.237251752 cg22496377 SHF S_Shore 5URT 39.0186 6.25E−10 0.453229504 0.598199089 cg22498143 SPT3N4 N_Shore Body 21.069 4.21E−06 0.640750977 0.792908022 cg22505086 DLX5 S_Shelf 3UTR 5.8853 0.000843 0.286864372 0.345839043 cg22511262 WT1 S_Shelf Body 9.41329 3.62E−07 0.159214555 0.274852851 cg22531668 VAX1 N_Shore 3UTR 14.584 6.99E−05 0.453199062 0.611852071 cg22833573 WT1 S_Shore TSS1500 13.7978 5.67E−08 0.14795094 0.250127828 cg22557091 PAX6 Island Body 14.6401 1.10E−09 0.158600792 0.309527235 cg22557662 PPP1R14A Island TSS200 19.236 1.08E−07 0.547188083 0.714845321 cg22559034 NR2F1-AS1 S_Shore Body 7.71168 1.24E−05 0.46278823 0.635338986 cg22560193 APC2 Island Body 10.4725 0.000593 0.399053443 0.474002812 cg22571664 NPR3 Island Body 9.92306 1.76E−09 0.21545106 0.423046241 cg22572908 ARHGEF4 S_Shore Body 6.76475 0.000913 0.280091768 0.362902791 cg22614142 KIFC2 Island Body 30.5204 4.24E−11 0.351119509 0.613047689 cg22616881 DLX6-AS1 N_Shelf Body 5.09098 0.001228 0.114079502 0.170060134 cg22623223 PTPRN2 Island Body 4.68816 0.000587 0.256707027 0.347335654 cg22637538 CHRNB1 Island TSS200 6.58602 4.36E−07 0.276723921 0.441389322 cg22660933 HOXB1 Island TSS200 5.04489 0.000476 0.55730887 0.635605899 cg22666373 OTX1 S_Shore Body 15.3527 3.38E−08 0.42074545 0.566590677 cg22674699 HOXD9 Island 1stExon 10.242 7.95E−06 0.059148891 0.124531201 cg22711111 FRIM15 Island Body 5.38684 9.93E−05 0.265351435 0.388795697 cg22720790 ZNF274 Island 5URT 9.41073 3.44E−05 0.562506674 0.667911899 cg22730140 NKX2-5 Island Body 10.0985 1.45E−06 0.124523364 0.213164766 cg22780464 GPR150 Island TSS200 10.7352 3.01E−07 0.258252384 0.403291076 cg22740492 PAX6 Island Body 17.759 1.59E−12 0.149710173 0.326517684 cg22770911 GATA3 S_Shore Body 13.7179 0.000541 0.577785783 0.665098929 cg22774038 JSRP1 Island Body 4.25009 1.37E−05 0.197505494 0.35245203 cg22777724 HOXB-AS1 S_Shore Body 3.55866 0.00072 0.357495003 0.467293199 cg22778788 NKX2-3 N_Shelf Body 6.64241 7.81E−05 0.342002399 0.435237929 cg22783363 TNFRSF10D Island TSS200 9.70479 8.00E−07 0.243123222 0.396629565 cg22799963 GATA4 Island 5URT 17.5528 6.93E−06 0.47390493 0.578669384 cg22807449 HOXB-AS1 S_Shore Body 4.52324 2.35E−05 0.320011555 0.455339277 cg22814146 BCAT1 N_Shore TSS200 14.4296 4.07E−06 0.158362062 0.280640049 cg22827250 PITX1 Island 3UTR 12.7205 2.34E−08 0.281253935 0.442493603 cg22839075 SYN2 N_Shore TSS1500 7.59931 0.000275 0.128141188 0.213019953 cg22840219 RCN3 Island Exon3nd 11.8477 1.42E−07 0.323252038 0.475138543 cg22849665 PRRT1 Island Body 9.19065 0.002107 0.703011247 0.776908717 cg22859061 SCGB3A1 Island TSS200 5.59467 7.02E−07 0.122388481 0.280112138 cg22859289 LOC101927248 S_Shore Body 4.75129 0.000621 0.276802955 0.360681745 cg22876812 LINC01143 Island Body 5.57656 1.47E−07 0.186988353 0.351329937 cg22878622 TCF24 Island Body 18.8152 1.43E−07 0.144264364 0.270953809 cg22882523 CPLAH Island Body 11.8791 6.70E−06 0.448950134 0.569208891 cg22888055 ITPKA S_Shore 3UTR 24.6165 7.52E−10 0.456490881 0.606547041 cg22897615 PRRT1 Island Body 3.84547 0.006841 0.519061517 0.697345682 cg22900229 POMC Island TSS1500 5.33411 0.000301 0.333085078 0.442527124 cg22902505 PRDM8 S_Shore 5URT 3.54023 0.00843 0.276370197 0.249599736 cg22937649 NTN1 Island Body 13.5905 1.68E−14 0.106284316 0.368860752 cg22951509 GPR150 Island TSS200 10.0424 1.67E−05 0.181613438 0.2762579 cg22973789 SATB2 Island 5URT 3.48414 0.00446 0.114521178 0.191954366 cg22975913 WT1-AS N_Shelf Body 3.80356 0.001312 0.351106634 0.452161401 cg22976224 SIM2 Island TSS1500 8.46301 7.98E−07 0.077695895 0.174327331 cg22982368 PAX6 Island TSS200 11.8471 1.29E−13 0.197057369 0.432168778 cg22997113 HOXA4 Island 1stExon 5.18912 0.006029 0.431223853 0.457729909 cg23016129 SPAG6 Island TSS200 21.0506 1.72E−12 0.178919671 0.364979757 cg23027574 RNF39 Island 1stExon 5.29116 5.34E−05 0.251970986 0.374812257 cg23042510 EBF3 Island Body 10.6104 5.61E−07 0.122017123 0.231835397 cg23044079 FOXI3 Island 1stExon 6.03851 0.000899 0.202877564 0.267768172 cg23051664 TNFRSF10D N_Shore Body 7.86272 8.87E−06 0.118847589 0.213830427 cg23054189 TRIM58 Island 1stExon 15.2492 3.88E−14 0.188259264 0.43685849 cg23064601 PITX1 Island Body 21.1745 3.81E−07 0.411121927 0.531312982 cg23068797 MIR199A1 TSS1500 20.7834 2.47E−05 0.671865202 0.797639919 cg23082339 PCDHGA12 S_Shelf TSS200 4.03889 0.002182 0.102595718 0.181629681 cg23097402 DMRTA2 Island 3UTR 5.48978 8.48E−05 0.216296137 0.327091954 cg23100152 AMPD3 Island TSS200 3.34808 0.00484 0.131480948 0.211547721 cg23104954 DLEU1 Island Body 5.41075 0.000286 0.42306875 0.522236282 cg23125492 PAX6 Island Body 15.9741 6.30E−11 0.251856022 0.429903922 cg23129930 HOXA-AS3 Island Body 17.4305 145E−07 0.338158326 0.487141859 cg23130254 HOXD12 Island 1stExon 13.3545 6.87E−08 0.124428058 0.22551666 cg23158731 COL14A1 Island 5URT 6.63561 1.36E−07 0.140430404 0.289315465 cg23165310 SOX2-OT Body 6.02918 2.83E−05 0.298211924 0.403143335 cg23167905 FGF19 S_Shore Body 3.50828 0.001774 0.198961576 0.262867646 cg23196831 COL14A1 Island 1stExon 11.5137 9.36E−08 0.191447871 0.520937458 cg23218559 AMH Island Body 12.4766 2.66E−09 0.232576255 0.405197975 cg23229261 OTX1 Island Body 7.10692 2.56E−09 0.165176781 0.355188044 cg23244790 PCDHGA12 Island 1stExon 8.30347 7.70E−10 0.231109879 0.45777518 cg23244913 HCG9 N_Shore Body 7.43065 2.46E−05 0.365494322 0.54155302 cg23268203 HORMAD2 Island TSS200 13.7572 2.32E−08 0.252420597 0.350956081 cg23278196 ATP5G2 Island TSS1500 3.87564 0.000705 0.038454962 0.186450567 cg23286645 SIM2 Island Body 4.86857 1.18E−05 0.219794027 0.354247731 cg23290344 NEFM Island TSS1500 12.4376 2.03E−14 0.139231716 0.372266651 cg23345004 KATNAL2 N_Shore TSS1500 7.96059 1.53E−05 0.500575095 0.659920881 cg23357257 SEZ6L2 Island Body 12.3914 2.73E−06 0.365022309 0.494450385 cg23442209 LEX1-AS1 N_Shore Body 6.5594 1.28E−05 0.196035255 0.324355519 cg23445461 PCDHGC4 Island TSS200 4.8621 2.15E−05 0.303451733 0.502209951 cg23483840 HLA-J Island Body 5.8642 4.60E−05 0.120279602 0.230990284 cg23497385 LOC151174 Island Body 6.4894 7.37E−06 0.341004812 0.497464582 cg23500724 RNF39 Island 1stExon 4.68294 0.000778 0.058372736 0.134162122 cg23501406 DRD4 Island TSS1500 26.0624 8.99E−11 0.245814052 0.417622791 cg23503176 LEX1-AS1 S_Shore Body 8.70704 5.13E−05 0.186153121 0.282366413 cg28211613 FOXI3 Island TSS1500 6.41775 3.90E−07 0.075169072 0.190512508 cg23513966 ISL2 Island Body 8.89981 0.000263 0.234522008 0.344287657 cg23514619 TTBK1 Island Body 4.02301 0.000856 0.125150023 0.205127556 cg23525400 TRIM71 Island TSS200 13.7847 7.19E−09 0.095515739 0.204913171 cg23530553 HOXB-AS1 S_Shore Body 4.96747 0.001176 0.470218775 0.559645081 cg23546474 PAX3 Island Body 5.39576 0.000368 0.463882674 0.552153538 cg23585322 COL14A1 S_Shelf 5URT 7.07347 1.43E−05 0.46711853 0.62595027 cg23587449 LRAT N_Shore 5URT 10.7775 2.50E−05 0.130083934 0.205671255 cg23598080 LHX4 Island Body 7.49566 0.000239 0.143535736 0.205230488 cg23519365 LINC00403 Island Body 8.63552 1.15E−07 0.103818198 0.221808533 cg23619399 RNF219-AS1 N_Shore Body 8.88903 1.65E−06 0.261460278 0.331275873 cg23623622 TRIM15 Island Body 4.46835 0.008133 0.151855743 0.217229153 cg23634124 DLX4 Island Body 4.92389 0.000471 0.257215491 0.350835693 cg23637494 LINC01391 N_Shore TSS1500 6.25804 4.67E−05 0.292538345 0.385950087 cg23655970 PAX9 S_Shore TSS200 4.41708 0.00161 0.163220498 0.231557932 cg23663774 PITX2 Island 5URT 9.30858 2.94E−08 0.067928751 0.164025673 cg23676151 NR2E1 Island Body 9.91836 1.13E−06 0.449251457 0.598177879 cg23683583 PAX9 Island Body 8.08678 7.29E−05 0.120659357 0.262138377 cg23695707 GSC Island 3UTR 4.62315 0.000142 0.168697961 0.281846693 cg23697546 HOXC6 S_Shore 3UTR 4.82912 0.000101 0.186174868 0.236545028 cg23707289 BLC2 S_Shelf TSS1500 8.45716 1.37E−05 0.218285002 0.319906794 cg23712018 RNF39 Island Body 10.1331 2.67E−07 0.140609447 0.257377802 cg23724641 LYPD1 Island 5URT 7.61791 0.000192 0.145698396 0.210498959 cg23740652 PTPRN2 S_Shore Body 7.13035 0.00589 0.803957512 0.859832646 cg23767994 DLX5-AS1 Body 7.90921 2.51E−06 0.331337767 0.448147059 cg23792314 BCAT1 Island TSS1500 12.6155 4.56E−08 0.140733788 0.258710807 cg23829024 HIST1H1A N_Shelf TSS200 7.68857 2.27E−07 0.229658204 0.380509071 cg23831143 TNFRSF10C Island 5URT 55.4052 8.36E−10 0.489023538 0.663491341 cg23847722 DRD5 Island TSS200 6.90811 1.47E−06 0.155066589 0.291915595 cg23874561 PRDMB Island Body 9.87044 6.78E−10 0.111619412 0.230528381 cg23880589 KCNAB3 S_Shore Body 29.753 1.05E−08 0.398107899 0.513836758 cg23896164 SIM2 Island 3UTR 21.0144 3.52E−06 0.510540417 0.613651929 cg23907108 NRXN2 N_Shelf Body 13.9937 1.05E−05 0.426980853 0.554542384 cg23917057 FOXI3 Island TSS1500 11.6 4.79E−09 0.220920102 0.373813957 cg23933618 DKFZp686K1684 Island TSS1500 6.36392 4.12E−07 0.157953229 0.303772567 cg23937893 WNT10A Island Body 6.23231 0.001129 0.264951309 0.355386735 cg23939808 RNF39 Island 1stExon 4.668 9.13E−05 0.019813884 0.198822377 cg23941075 PCDHB15 S_Shelf 5URT 9.35336 5.48E−05 0.203467116 0.309065099 cg23965061 TNFRSF10C Island 5URT 12.2175 5.39E−08 0.178507434 0.318997626 cg23971069 PROX1-AS1 N_Shelf Body 19.9112 1.20E−07 0.267771527 0.36612499 cg23973429 GRHL2 Island 5URT 85.3366 7.33E−09 0.407950783 0.566660424 cg23974473 PCDHB15 N_Shore 1stExon 10.4647 5.39E−05 0.617862039 0.727866848 cg23979631 HOXA2 N_Shelf TSS200 7.47565 1.10E−05 0.173805747 0.272675545 cg23994043 SATB2 Island 5URT 4.47791 0.000233 0.273602127 0.361899128 cg24001710 EN1 Island TSS1500 5.04029 0.000374 0.175106502 0.246951919 cg24005685 PITX2 Island Body 14.273 3.52E−10 0.157821613 0.272116806 cg24008908 AKR7A3 Island TSS200 12.6395 3.09E−08 0.366760932 0.568626313 cg24016627 RNF39 Island 1stExon 4.3238 6.15E−05 0.081081917 0.194659067 cg24030449 FGF20 S_Shore 1stExon 5.23328 0.000118 0.412238089 0.552448481 cg24059697 GATA3-AS1 Island Body 3.43044 0.000238 0.245337404 0.409261579 cg24051554 C2CD4D Island 5URT 7.34634 4.93E−07 0.285581122 0.447180062 cg24080247 SIM2 S_Shore Body 6.72944 6.92E−07 0.161306786 0.295104493 cg24157892 T8X2-AS1 Island Body 5.88056 3.92E−06 0.119245805 0.25055837 cg24169669 TTC6 Island Body 8.14047 7.17E−11 0.219806976 0.452130458 cg24225707 DLX1 S_Shelf TSS1500 5.68754 3.79E−05 0.2319201 0.890805525 cg24241323 HOXB3 S_Shelf 5URT 3.53374 0.009604 0.590567566 0.659156768 cg24280332 C10orf11 N_Shore Body 9.31918 0.001074 0.76451676 0.854504778 cg24309555 APOB Island TSS200 6.81403 0.000727 0.47869114 0.574216768 cg24328539 GATM Island TSS200 5.46503 0.007076 0.240330861 0.331177869 cg24330456 RNF39 N_Shelf Body 10.5698 1.53E−05 0.662954544 0.801047505 cg24351901 HLA-F Island Body 31.2631 1.57E−07 0.466853888 0.583886788 cg24354581 SIM2 N_Shore Body 4.58415 0.000412 0.213289015 0.305992435 cg24370475 FGF19 S_Shore Body 5.74453 1.92E−05 0.142829115 0.244095591 cg24375215 CD3COA 5URT 8.45514 8.18E−07 0.314102137 0.464290485 cg24382527 ADAM8 Island 1stExon 4.31291 0.000739 0.124411855 0.214731954 cg24425021 POU451 N_Shelf 1stExon 13.3206 8.50E−09 0.05114992 0.159439107 cg24425171 POMC Island TSS200 5.51594 0.000516 0.450558334 0.585620717 cg24425838 C2CD4D Island 5URT 8.04978 4.06E−05 0.577504616 0.716433145 cg24426072 TRIL Island Body 9.10441 3.87E−07 0.096674815 0.19428478 cg24430580 PITX2 Island 1stExon 18.593 2.33E−10 0.200261234 0.335632042 cg24446586 HOXA11-A5 Island Body 18.7192 8.44E−07 0.419317947 0.538348879 cg24453580 PRPH Island Body 4.08088 0.000155 0.1640223 0.272120074 cg24479590 CHRN81 Island TSS200 6.48538 7.18E−07 0.243173068 0.400215708 cg24524285 NRXN2 N_Shore Body 12.672 8.09E−07 0.479458597 0.602308899 cg24585439 MMP235 Island TSS1500 16.759 3.48E−12 0.268441284 0.480289835 cg24550149 TTC22 Island Body 8.97432 1.94E−05 0.563631047 0.674299828 cg24565496 LMF1 Island TSS1500 58.2216 3.20E−012 0.327989706 0.491670521 cg24575083 CYBSR2 Island TSS1500 6.53952 1.27E−05 0.332238009 0.473399612 cg24582934 RGCC S_Shore Body 6.77881 3.73E−05 0.23754295 0.358438555 cg24592027 DBX1 N_Shore TSS1500 5.67504 0.001973 0.14378014 0.197944163 cg24628744 H2AFY S_Shore TSS1500 7.57965 4.39E−07 0.298722368 0.430587136 cg24630419 ESRRG Island TSS1500 9.71741 1.23E−06 0.225442794 0.340091548 cg24633978 HOXD11 S_Shore 1stExon 14.6389 7.33E−10 0.13621414 0.286895208 cg24646414 GATA4 Island 5URT 8.6012 8.40E−07 0.274792632 0.414211108 cg24657817 BEND4 Island Body 11.7396 2.54E−11 0.13015339 0.29672914 cg24663845 SLC4A15 TSS200 22.9315 4.13E−05 0.491421442 0.628085667 cg24663414 GFI1 Island TSS1500 14.7337 7.29E−05 0.504974193 0.597381488 cg24697184 TTC22 Island 1stExon 12.7557 5.48E−07 0.329935793 0.458033191 cg24701575 DKFZp686K1684 N_Shore Body 10.6083 1.58E−08 0.349858945 0.532733919 cg24721899 NKX3-2 Island Body 12.1962 9.34E−12 0.151703874 0.33521706 cg24725574 HLA-J Island Body 5.07393 0.000682 0.07796217 0.15459086 cg24727395 ORFPR Island TSS200 7.65875 3.66E−08 0.113600428 0.272340194 cg24745495 EPHX3 Island TSS1500 71.2682 6.88E−15 0.293213641 0.468856086 cg24747764 LINC00403 S_Shore TSS1500 9.47399 9.84E−04 0.155334642 0.264396424 cg24787138 TTBK1 Island Body 11.0163 1.06E−05 0.44878814 0.587278929 cg24796272 STK3 N_Shore 1stExon 4.51356 0.005053 0.364208086 0.433055798 cg24797840 GATA3-AS1 Island Body 3.08176 0.006193 0.244332112 0.33097299 cg24804179 PRPH Island Body 3.92504 0.008599 0.186555185 0.251161328 cg24880701 SKOR1 Island Body 12.0383 1.40E−09 0.18221125 0.393152841 cg24896649 ZFP42 Island TS200 12.5101 1.40E−13 0.162287041 0.92712148 cg24901042 TMPRSS2 Island TSS1500 3.83391 0.009399 0.290989035 0.359891185 cg24918868 NKX2-3 N_Shelf 5UTR 10.0149 1.10E−06 0.314263032 0.412929301 cg24914355 HOXD13 S_Shelf Body 11.7194 1.64E−09 0.204600198 0.380250256 cg24927800 DES Island 1stExon 10.5912 1.35E−10 0.140562602 0.326382139 cg24928391 SOX17 Island TSS200 5.23559 0.000496 0.081429571 0.16216134 cg24933175 SHF S_Shore 5URT 7.38014 7.66E−06 0.094971677 0.19939695 cg24951286 PCDHB15 S_Shelf 5URT 7.74918 4.36E−05 0.290059291 0.37688902 cg24958321 TRIM71 S_Shore Body 4.69296 0.00569 0.381839648 0.456402129 cg24988255 HOXA11-AS Island Body 5.53106 0.000533 0.265744748 0.340130847 cg25005858 SPEG Island Body 21.5039 3.99E−06 0.504573092 0.624879778 cg25032094 MYO158 Island TSS1500 15.331 1.08E−07 0.456509021 0.620584928 cg25034395 HPSE2 S_Shelf TSS1500 5.09092 0.000662 0.402043378 0.514921875 cg25035485 APOB N_Shore Body 3.34154 0.005895 0.477120519 0.585136599 cg25044651 LVRN Island 1stExon 12.6562 2.05E−07 0.209436771 0.332881008 cg25051331 LOC101929154 Island TSS1500 7.32715 0.000242 0.177458389 0.243695189 cg25062797 VAX1 Island Body 3.51017 0.003409 0.18238856 0.272392099 cg25071744 APCB S_Shore TSS200 8.99596 0.001005 0.560700209 0.637106313 cg25083618 CYP26C1 Island TSS200 7.0896 0.001695 0.609983659 0.685111354 cg25112812 ZNF518B Island TSS1500 14.871 4.15E−07 0.288440614 0.395711422 cg25125060 NR2E1 Island TSS1500 7.42331 0.001244 0.482871279 0.552940444 cg25160978 GDF7 N_Shelf TSS1500 7.65188 9.43E−07 0.10427144 0.281791465 cg25161512 PON3 Island TSS1500 11.1007 1.25E−06 0.261804003 0.375530995 cg25188395 HOXA9 Island Body 4.37431 0.000316 0.171249677 0.259552867 cg25192855 IRF8 N_Shore 5URT 16.6074 2.95E−09 0.421099306 0.6404799 cg25196158 PROX1-AS1 N_Shelf Body 4.87306 0.000105 0.250761262 0.35518323 cg25200152 NBPFS S_Shore Body 6.97059 3.27E−07 0.233071439 0.378569403 cg25203481 KCNQ1DN Island TSS200 5.19083 8.70E−05 0.260605263 0.378628758 cg25212090 LHX9 S_Shore Body 7.49612 7.25E−05 0.167339743 0.254862429 cg25247290 WT1-AS S_Shore Body 16.7301 1.58E−13 0.196596173 0.372124214 cg25258843 SOX7 Island Body 6.04651 0.000114 0.363064993 0.479177468 cg25307665 HOXA5 Island TSS1500 5.93145 4.11E−05 0.324350438 0.42384864 cg25318809 HLA-1 Island Body 5.64026 2.09E−05 0.113565088 0.227900399 cg25330243 GRHL2 Island TSS200 8.01586 8.42E−09 0.189124865 0.373253655 cg25336332 THR5-AS1 Island Body 4.89357 0.000386 0.556342095 0.67117136 cg25342207 EBF3 Body 18.1495 0.001497 0.747840395 0.810058859 cg25371919 OTX1 S_Shore Body 10.513 4.56E−07 0.488625012 0.634837316 cg25377206 TBX20 Island TSS1500 14.328 9.57E−09 0.413568473 0.349091909 cg25382400 SOX2-OT S_Shore Body 7.50343 1.84E−05 0.269019012 0.387940161 cg25390165 HOXA5 Island TSS1500 11.6613 0.000146 0.397156882 0.458243424 cg25412546 OTP Island 3UTR 9.62002 1.94E−06 0.168593493 0.261996869 cg25437411 HPSE2 S_Shelf TSS1500 3.27338 0.007114 0.153940144 0.230454038 cg25442600 IRX4 S_Shore TSS1500 3.7993 0.001734 0.412884144 0.512836301 cg25471923 HOTTIP N_Shelf Body 3.16751 6.87E−05 0.304269897 0.406051929 cg25478600 KCNO1DN Island TSS1500 4.39621 0.000306 0.222245894 0.328805502 cg25479916 HIST1H31 S_Shore TSS1500 7.44871 8.35E−08 0.223337767 0.373628442 cg25485294 HIST1H31 S_Shore TSS1500 8.39554 6.04E−09 0.20296936 0.388257668 cg25506432 HOXA5 Island TSS1500 10.37 0.000356 0.671558364 0.751274385 cg25527090 VAX1 Island Body 4.75398 0.000603 0.22171125 0.315296307 cg25538137 GATAS-AS1 Island Body 5.69024 0.000979 0.140877152 0.000072837 cg25539045 FOXI2 Island TSS1500 21.2076 2.42E−07 0.435622679 0.617786807 cg25547580 ONECUT2 Island 1stExon 8.44103 1.03E−09 0.124406002 0.291281792 cg25576011 FOXI2 Island TSS200 7.79966 1.66E−07 0.173596996 0.332067053 cg25586361 WT1 N_Shelf Body 5.1879 0.001054 0.34646815 0.431798869 cg25593794 TP73 5URT 9.26235 0.003298 0.659005739 0.706849687 cg25608490 WNT16 N_Shore Body 9.8664 1.54E−06 0.320328894 0.451435535 cg25622366 OTX1 Island Body 13.1368 3.37E−15 0.153557254 0.408855264 cg25639415 GRHL2 Island TSS200 8.46181 2.24E−07 0.226757884 0.390591678 cg25678088 DLX6-AS1 S_Shore Body 15.713 1.70E−07 0.307650186 0.414402939 cg25685262 FDX1 S_Shore TSS1500 17.3908 3.58E−07 0.19105396 0.280696765 cg25702780 MECOM N_Shelf 5URT 7.75589 0.000101 0.099598867 0.180158912 cg25720804 TLX3 Island 1stExon 11.22 2.70E−10 0.114630116 0.265155621 cg25746778 EBF3 Island Body 3.84577 8.24E−05 0.390170692 0.561990607 cg25755575 ESRRG Island TSS1500 10.5222 9.44E−09 0.186347842 0.333520779 cg25756435 MAST1 Island Body 14.1342 1.12E−10 0.056817843 0.193368065 cg25775494 NR2E1 S_Shore Body 12.4111 1.04E−08 0.192080868 0.315540502 cg25782229 WT1 N_Shore Body 5.08188 5.53E−05 0.291352308 0.426482596 cg25803093 SPAGS Island TSS1500 12.0008 4.58E−07 0.261862001 0.382851879 cg25835733 GAD2 S_Shelf TSS1500 16.7435 3.21E−06 0.40886986 0.51689504 cg25845597 HIST1H51 Island TSS1500 22.2872 1.37E−11 0.102347663 0.228474179 cg25845985 PRRT1 Island Body 7.58947 0.00111 0.398367349 0.479750438 cg25851789 ESRRG N_Shore 5URT 7.55386 1.18E−07 0.238501846 0.39876221 cg25884711 NPY Island 1stExon 6.57281 6.22E−07 0.067096877 0.173597491 cg25887069 GALRS Island Body 8.14395 7.99E−06 0.074521401 0.1795653 cg25903779 LOC101929076 Island Body 4.43091 2.06E−05 0.239491807 0.387931609 cg25903783 TBX2-AS1 Island Body 7.76112 0.000577 0.412628961 0.489641374 cg25916711 NKX2-5 Island 3UTR 11.4637 4.57E−09 0.108888036 0.242450121 cg25923240 SOX27 S_Shore 3UTR 4.38709 0.004845 0.511143581 0.538707525 cg25930229 LBX1-AS1 S_Shore Body 13.9234 8.77E−08 0.451616922 0.625450517 cg25939676 SHISA5 N_Shore TSS1500 7.39368 9.41E−06 0.117279008 0.207519362 cg25942450 TLX3 Island TSS200 10.9743 1.08E−06 0.324578629 0.460319566 cg25960088 HOX5-AS3 N_Shore Body 9.70669 1.87E−07 0.410568783 0.561982182 cg25965471 TTBK1 Island 1stExon 5.93549 1.40E−05 0.336181665 0.468041207 cg25984671 TCF15 Island TSS200 8.43721 2.22E−08 0.134977915 0.289996672 cg25988034 GATA5-AS1 Island Exon3nd 4.38813 0.000249 0.192513788 0.306498474 cg28000619 AMH Island Body 7.39291 4.48E−08 0.348938591 0.548758398 cg26006870 GATA4 S_Shore 5URT 6.81839 0.000154 0.42045824 0.520033808 cg26006951 TERT S_Shore TSS1500 18.8185 3.42E−06 0.331547036 0.423735485 cg26019112 PAX6 Island Body 21.2925 2.34E−12 0.188593009 0.358015893 cg26057751 CDSA Island Body 8.88691 2.27E−07 0.292058771 0.445992855 cg26058714 MECOM Island TSS200 7.91242 0.000188 0.172968013 0.26075307 cg26060971 DNAH1 Body 14.689 0.000424 0.599878248 0.636936646 cg26067760 HPSE2 S_Shelf TSS1500 7.38632 5.07E−07 0.375213837 0.525892972 cg26069745 HOXA2 N_Shelf 1stExon 18.0658 3.34E−06 0.351834764 0.44697104 cg26072906 BATF3 S_Shore TSS1500 6.68444 1.06E−07 0.17365335 0.362048732 cg26094599 WNT10A Island Body 76.9584 2.04E−07 0.591748767 0.700005727 cg26094789 PAX9 Island Body 5.64759 0.000592 0.435690047 0.344594788 cg26099134 SIM1 Island Exon3nd 7.58624 7.43E−06 0.268121495 0.371403074 cg26125683 FOXI2 Island 1stExon 8.05555 8.71E−08 0.38540356 0.558311436 cg26144458 SIM2 Island TSS1500 8.72753 5.94E−05 0.261333946 0.351203622 cg26149275 EVX2 Island TSS1500 7.15986 4.85E−07 0.236275136 0.399529956 cg26151467 SOX1 S_Shore 3UTR 4.33594 0.000215 0.241894916 0.357166732 cg26160218 THR5-AS1 S_Shore Body 4.27155 0.00289 0.569530752 0.670255192 cg26176204 AGAP1 N_Shore TSS1500 13.713 8.15E−08 0.326480151 0.483341919 cg26190890 HIST1H2BK S_Shore 3UTR 5.04842 0.000278 0.484847692 0.604845826 cg26195178 IRX4 Island 5URT 4.08092 0.000317 0.134972206 0.235357774 cg26199906 OPLAH Island Body 6.909 3.37E−05 0.297374112 0.412710604 cg26206945 NKX2-5 Island TSS1500 26.0679 1.73E−09 0.385687357 0.527252323 cg26212180 VAX1 S_Shore TSS1500 8.46876 1.54E−06 0.151715372 0.255673904 cg26214742 H2AFY S_Shore TSS1500 16.5714 6.16E−08 0.467164116 0.586909495 cg26224060 GPR150 Island 1stExon 17.3302 1.80E−08 0.302262295 0.422238172 cg26241806 APB31IF Island TSS200 9.77631 2.78E−07 0.099475271 0.205115261 cg26248075 SIM1 S_Shore Body 6.28913 4.92E−05 0.385692 0.4846957 cg26254045 SPRED3 S_Shore Body 7.04285 0.002009 0.52900457 0.801639822 cg26256263 RUNX3 Island TSS1500 5.70081 0.00062 0.348527662 0.449295081 cg26259537 HOXA10-AS Island Body 6.73685 5.23E−05 0.155708135 0.254192375 cg26261502 DKFZp686K1684 N_Shore Body 8.75406 8.13E−06 0.184531733 0.26259027 cg26263263 VAX1 S_Shore Body 6.72144 0.000285 0.304858898 0.386230222 cg26277730 BCL2 S_Shelf TSS1500 4.63522 7.95E−05 0.191822874 0.308971636 cg26277754 LOC642366 Body 11.6833 2.75E−09 0.183016834 0.333655734 cg26295921 PTPRN2 N_Shore Body 11.7045 0.000133 0.598315244 0.631481646 cg26295488 DRDS Island TSS200 4.13921 6.41E−05 0.149136715 0.281459119 cg26299189 PDX1 Island 3UTR 4.77207 9.52E−05 0.114245238 0.216815945 cg26309194 TMPRSS2 Island TSS1500 4.33724 0.001059 0.384399108 0.483950244 cg26322248 ISM1 Island TSS1500 12.7477 1.43E−08 0.078598664 0.20826524 cg26827071 HOXB4 S_Shore TSS200 9.37065 5.56E−07 0.109497328 0.198771454 cg26330518 NEFM N_Shore TSS1500 17.2163 5.80E−10 0.807635028 0.508179908 cg26531172 SEMA6C Island 5URT 5.11016 2.93E−05 0.349950231 0.518152009 cg26565545 OTP N_Shelf Body 7.9715 5.46E−07 0.301894073 0.447436076 cg26870729 F2RL1 Island Body 6.90473 2.85E−07 0.139148714 0.296043 cg26371731 C1orf61 N_Shore 5URT 6.76108 2.40E−07 0.216201957 0.365090393 cg26373163 IERS Island TSS200 5.00339 7.94E−04 0.22004826 0.36022581 cg26380443 PCDHB15 S_Shelf 1stExon 3.80615 0.003727 0.142221797 0.213678939 cg26388730 NRXN2 Island Body 7.24742 0.003485 0.535759465 0.600493909 cg26410485 LHX4 Island Body 5.44455 0.000262 0.074242948 0.14889501 cg26413174 TTC22 Island 1stExon 6.65187 6.30E−05 0.185177553 0.268575959 cg26421310 RUNX3 Island TSS1500 4.47804 0.004434 0.137557845 0.20383534 cg26422458 ADGRL4 Island 1stExon 8.0834 3.99E−08 0.211370453 0.383782913 cg26429925 BNC1 Island TSS1500 5.1253 0.0009 0.498803319 0.603335596 cg26437825 LOC648987 S_Shore TSS1500 6.81687 2.85E−06 0.206528777 0.351434737 cg26446499 LOC101929154 N_Shore TSS1500 12.0122 2.98E−12 0.239723033 0.455454442 cg25457809 KCNO1DN Island TSS1500 5.69035 0.000164 0.36545956 0.485924214 cg26476852 HCXA8 Island 1stExon 13.5927 3.67E−07 0.176670594 0.270323785 cg26506288 HOXB13 S_Shelf 3UTR 5.19516 8.18E−05 0.217535988 0.831205621 cg26417171 TLX3 Island TSS1500 8.87594 2.60E−07 0.210043347 0.324803151 cg25517714 TLX3 Island TSS200 16.0372 5.44E−09 0.186080333 0.302858773 cg26521404 HOXA9 Island 1stExon 11.9081 2.05E−08 0.066737467 0.172729722 cg26551975 HOXA10 N_Shelf Body 4.10625 0.000433 0.159896935 0.259631492 cg26559315 LYNX1 Island 5URT 4.97513 5.20E−06 0.160019299 0.295721502 cg26560222 DMRTA2 Island 1stExon 8.18753 5.07E−07 0.224504707 0.332990389 cg26585452 DLEU1 S_Shore Body 8.03856 0.000822 0.707903992 0.773862505 cg26589785 ITPKA Island Body 18.5156 1.43E−08 0.366107456 0.529562714 cg28601317 KCNQ1DN Island TSS1500 4.74595 0.001343 0.278491437 0.359453891 cg26618965 TCF24 Island 5URT 14.0539 1.56E−08 0.087905065 0.196064312 cg26559605 AMH Island Body 13.4807 1.84E−09 0.279336936 0.497531811 cg26560900 TBX2 Island Body 5.76442 0.001105 0.458101093 0.582014141 cg25573012 TBX20 Island TSS200 33.7769 0.01E−15 0.070220453 0.192037793 cg26674479 EVX1 N_Shore TSS1500 8.48421 0.000201 0.421966543 0.515589576 cg26674943 ISL2 N_Shore TSS1500 6.40645 0.003825 0.12725121 0.178737921 cg26687072 HOXB1 Island TSS200 5.97237 0.000281 0.429621984 0.514800884 cg26690075 WNT16 N_Shore Body 11.7612 8.25E−09 0.333750395 0.508539336 cg26698460 ZNF274 Island 5URT 9.32518 0.000159 0.56225555 0.640661071 cg26701242 SIM2 N_Shore Body 7.32496 3.72E−05 0.219306933 0.295784498 cg26702958 LBX1-AS1 Island Body 10.6897 6.32E−09 0.240206662 0.414780264 cg26708319 PITX2 N_Shore Body 7.25162 1.99E−08 0.118863698 0.30390468 cg26751356 LINC00403 Island Body 10.4959 1.57E−09 0.158902635 0.295522499 cg26755097 FLOT1 Island TSS1500 10.3497 5.14E−10 0.210337051 0.39845602 cg26783087 VAX1 N_Shore 3UTR 7.50764 4.30E−08 0.200365347 0.352009825 cg26796135 GATM Island TSS200 3.97259 0.007881 0.195812144 0.277825016 cg26821372 PCDHGA8 N_Shore 5URT 4.94074 0.000438 0.427603122 0.519320922 cg26814075 LEP Island TSS200 39.8104 6.47E−11 0.304399043 0.422158867 cg26816907 LHX9 Island Body 11.7079 2.94E−07 0.34004184 0.474208497 cg26844246 TLX8 Island TSS200 14.7228 6.07E−10 0.236601845 0.401345928 cg26848718 WT1 S_Shelf Body 25.0638 6.59E−15 0.170813747 0.372272164 cg26870725 MAST1 S_Shore ExonEnd 9.22225 1.80E−08 0.162565004 0.818644245 cg26879945 PLLP Island TSS1500 8.54738 8.00E−05 0.253532887 0.36326841 cg26890354 PCDHGCA N_Shore TSS1500 6.04397 1.47E−05 0.228546397 0.341574774 cg26951440 LINC01475 S_Shore TSS1500 8.08893 1.62E−05 0.21718056 0.31402834 cg26990102 EN1 Island TSS1500 5.70395 1.84E−06 0.1999159 0.317105949 cg26993537 OTX2 S_Shore TSS200 5.54397 0.002488 0.315328522 0.398058144 cg27009763 HOXA8 N_Shelf 1stExon 6.46776 0.000188 0.278100441 0.374885937 cg27011060 PAX6 N_Shore 5URT 17.5783 2.29E−09 0.313376714 0.498753046 cg27029321 PRDM14 Island TSS200 14.4148 1.42E−05 0.32864667 0.418247576 cg27032146 DLX5 Island Body 3.82544 0.002069 0.264300564 0.361466154 cg27035251 JAK3 S_Shore 1stExon 11.6696 0.000194 0.332023276 0.423882606 cg27076669 CPM Island TSS1500 12.291 2.98E−07 0.225567825 0.32843323 cg27090216 TNFRSF10C Island 5URT 45.8607 3.07E−11 0.366740871 0.533705566 cg27100227 EVX1 S_Shelf TSS1500 3.80347 0.000844 0.257902582 0.360956638 cg27110277 FGF20 S_Shore TSS1500 14.9989 0.000127 0.560840682 0.758241894 cg27121250 PRDM8 S_Shore 5URT 5.12804 0.002436 0.384845171 0.44477299 cg27121463 FOXI3 Island Body 13.1549 2.17E−10 0.073416725 0.221890714 cg27125849 TBX2-AS1 Island Body 6.58446 6.62E−05 0.180957402 0.227827024 cg27151903 HOXA-A53 Island Body 4.91726 0.001416 0.332094095 0.426663074 cg27157482 HOXA10-HOXA9 N_Shelf Body 7.84846 4.90E−05 0.494402574 0.586097909 cg27162455 KCNA83 N_Shelf TSS1500 12.5188 6.88E−05 0.640068473 0.731904596 cg27183188 ALDOC TSS200 12.1761 8.00E−08 0.272570781 0.459714938 cg27190967 SYNGR8 N_Shore TSS1500 6.01813 0.000215 0.422827019 0.531480815 cg27234067 WT1-A5 Island Body 7.93953 1.09E−05 0.260355707 0.362308911 cg27252696 SIM1 Island TSS1500 9.00942 7.13E−09 0.112519336 0.253240779 cg27267227 SEZ6L2 Island Body 5.03417 6.05E−06 0.303641527 0.459877785 cg27304406 BNC1 Island TSS1500 8.36286 7.75E−05 0.324330193 0.416052447 cg27309110 JAK3 Island ExonEnd 4.78704 0.0095 0.526012488 0.564694414 cg27312652 SIM2 Island Body 10.2251 3.32E−05 0.176516904 0.235390672 cg27363141 RNF89 N_Shelf Body 66.564 0.000183 0.840916941 0.915282121 cg27364741 OTX1 Island Body 9.5317 3.67E−10 0.261176026 0.453866912 cg27398720 PITX2 S_Shore Body 8.55604 0.000522 0.375857775 0.442952053 cg27448110 PTPRN2 Island Body 20.5199 3.77E−07 0.354946636 0.465067303 cg27449131 IER5 Island Body 11.9179 1.24E−11 0.161100357 0.371042507 cg27493649 RNF39 Island Body 5.51227 0.000122 0.207871899 0.311842277 cg27513574 SLC34A2 Island 1stExon 5.86041 1.43E−06 0.180614395 0.328174817 cg27520775 ESRRG S_Shore TSS1500 4.69703 6.98E−06 0.195413895 0.828023662 cg27521476 TLX1 Island Body 4.31476 0.000348 0.339844224 0.451746313 cg27526665 THRB-AS1 Island Body 6.427 1.03E−05 0.233645664 0.347253078 cg27532187 RNF39 Island 1stExon 6.99183 2.17E−05 0.303351074 0.432222583 cg27539480 HOXA3 S_Shore 3UTR 7.1808 0.003141 0.746917287 0.812943484 cg27542552 ACF5 Island 5URT 10.0557 1.35E−05 0.176032701 0.266421849 cg27583307 SAT62 Island Body 4.24446 0.000343 0.453051023 0.596278901 cg27616661 SYNGR3 S_shore Body 4.50082 0.002228 0.070548111 0.14231126 cg27622506 THRB-AS1 Island Body 4.16577 0.000678 0.271823889 0.867695458 cg27639457 CTNND2 Island Body 13.7883 3.50E−05 0.344498339 0.426716335 cg27639662 PRDM3 S_Shore 5URT 12.1934 1.67E−05 0.378570512 0.451179465 cg27650870 ANK1 Island Body 5.20915 0.004457 0.40599055 0.469508383 cg27656658 HOXB-AS1 N_Shelf Body 3.97558 0.009116 0.418402057 0.450684586 cg27661315 BEND4 Body 6.36107 9.38E−05 0.449504772 0.552114101 cg12627726 KCNQ4 Body 0.020674 1.92E−09 0.689301388 0.583260727 cg06749480 PBX1 Body 0.104414 5.90E−06 0.779912519 0.689612949 cg07152605 PBX1 3UTR 0.077624 2.88E−07 0.709354581 0.607090081 cg12689805 TRIM58 3UTR 0.106512 2.16E−05 0.833537194 0.745223758 cg22029015 AGAP1 S_Shore Body 0.48735 1.84E−17 0.907829836 0.70669701 cg10185395 AGAP1 S_Shore Body 0.017036 1.33E−17 0.888728558 0.751125212 cg02408098 TRIM71 Body 0.092085 5.08E−07 0.856285419 0.760925616 cg20789595 ADCY5 Body 0.052227 2.57E−10 0.737646535 0.628378657 cg19008133 ADCY5 Body 0.072667 7.89E−05 0.59904531 0.540487677 cg07366462 ADCY5 N_Shelf Body 0.042033 5.68E−08 0.763565481 0.680088404 cg01246398 ADCY5 N_Shore Body 0.008935 2.48E−07 0.40769562 0.313433704 cg06598988 ADCY5 Island 1stExon 0.031784 5.19E−11 0.705115095 0.531770202 cg25661981 ADCY5 Island 1stExon 0.099409 4.44E−10 0.788356822 0.587644677 cg22989209 TERT N_Shelf Body 0.058527 4.58E−07 0.727614891 0.630747152 cg04830757 LINCC0461 Body 0.131857 7.67E−05 0.717647845 0.635406434 cg25863842 DDX39B N_Shelf Body 0.134966 0.002986 0.717371922 0.673810131 cg26177760 DDX39B N_Shore Body 0.154181 0.003356 0.552406372 0.499392 cg24124954 DDX39B N_Shore ExonEnd 0.085152 5.67E−05 0.537161775 0.459137556 cg24332685 DDX39B N_Shore Body 0.155806 0.000119 0.518894326 0.428966456 cg21179831 DDX39B N_Shore Body 0.098026 9.45E−05 0.528587674 0.459850232 cg21079591 DDX39B S_Shore TSS1500 0.029469 6.20E−11 0.5508733 0.971283727 cg24711558 CUX1 Body 0.13345 2.39E−05 0.64200962 0.54130583 cg03787603 CUX1 Body 0.107143 2.32E−08 0.897665512 0.776925732 cg11911769 CUX1 Body 0.06142 1.45E−09 0.740554333 0.621220253 cg09244071 CUX1 Body 0.068615 3.76E−08 0.712450031 0.605724939 cg15755348 CUX1 Body 0.140885 7.81E−06 0.801083481 0.697959239 cg26651148 PTPRN2 Body 0.176769 8.00E−05 0.817612767 0.721452768 cg14759472 PTPRN2 Body 0.155188 1.63E−06 0.907684791 0.762914495 cg26170014 PTPRN2 Body 0.129983 1.45E−06 0.826677845 0.710134414 cg10473311 PTPRN2 Body 0.082167 4.32E−06 0.593989473 0.485019758 cg21584493 PTPRN2 Island Body 0.046209 7.20E−08 0.641590783 0.558389717 cg26793019 CSMD1 3UTR 0.044095 1.01E−10 0.875629806 0.769001051 cg02563011 CSMD1 Body 0.121439 1.57E−05 0.754183341 0.659321232 cg02616558 CSMD1 Body 0.077825 2.84E−06 0.694586605 0.604867343 cg22663467 CSMD1 Body 0.098811 0.000222 0.825979853 0.769627687 cg26837853 CSMD1 Body 0.072424 1.00E−06 0.777059845 0.647951334 cg12269374 CSMD1 Body 0.057368 3.18E−09 0.725910907 0.600711755 cg08235285 CSMD1 Body 0.08072 4.55E−11 0.74507245 0.606898727 cg12042060 CSMD1 N_Shelf Body 0.050705 5.30E−11 0.772679473 0.634973586 cg14207559 C10orf11 Body 0.056885 7.75E−12 0.750300574 0.544735707 cg23264016 C10orf11 Body 0.032029 1.94E−11 0.672926256 0.480734505 cg26193427 INPP5A N_Shore Body 0.262155 0.003355 0.457120976 0.370231695 cg12507669 INPP5A N_Shore Body 0.17177 0.001573 0.420868841 0.888358101 cg05714496 INPP5A N_Shore Body 0.199455 0.000125 0.687561442 0.549142131 cg22900057 INPP5A S_Shelf Body 0.138777 6.81E−07 0.864730078 0.761965776 cg09746085 INPP5A Body 0.028091 1.28E−13 0.825529953 0.712523697 cg00221747 IGF2 N_Shore 3UTR 0.091459 7.06E−05 0.911026109 0.85081403 cg05334664 IGF2 S_Shore Body 0.207634 0.009981 0.780879988 0.700481899 cg06029905 IGF2 N_Shelf Body 0.062536 4.00E−07 0.742179512 0.656060717 cg02808220 IGF2 N_Shelf Body 0.080443 3.05E−05 0.781334566 0.719292525 cg24183187 IGF2 S_Shelf Body 0.12449 0.000146 0.760037225 0.66759202 cg22932998 IGF2 S_Shelf Body 0.061998 7.37E−07 0.838422411 0.764550616 cg21728792 IGF2 S_Shelf Body 0.089482 1.05E−05 0.806911209 0.723529788 cg24890224 IGF2 S_Shelf Body 0.061468 1.65E−06 0.604576371 0.516556253 cg01668279 IGF2 S_Shelf Body 0.098835 8.93E−06 0.799120567 0.710336727 cg16794682 CCND1 S_Shelf Body 0.024588 1.54E−12 0.789872977 0.666508789 cg12266049 CCND1 Island Body 0.052248 3.06E−11 0.913315481 0.77688797 cg15974867 CCND1 S_Shelf Body 0.11128 1.46E−10 0.833008631 0.651952645 cg04717045 CCND1 N_Shore 3UTR 0.018865 5.31E−12 0.688396899 0.546288869 cg12263469 CCND1 Ialand 3UTR 0.016578 5.73E−11 0.744055566 0.634779455 cg05569955 ATP5G2 N_Shore Body 0.126606 9.60E−05 0.478414854 0.374700999 cg26013733 ABR S_Shelf Body 0.054933 1.22E−08 0.744664636 0.634846232 cg13876604 ABR N_Shelf Body 0.122064 5.85E−07 0.739549372 0.615556829 cg08458121 ABR Island Body 0.20146 0.001866 0.862431147 0.803541707 cg02060732 RBFOX3 5URT 0.04139 2.10E−09 0.933731442 0.714892823 cg05604487 RBFOX3 5URT 0.077287 1.80E−06 0.657583581 0.545713354 cg04115702 RBFOX3 5URT 0.102341 3.31E−05 0.610661302 0.534299768 cg07490808 GREB1L Body 0.03196 1.00E−08 0.503961326 0.368909526

TABLE 10 Gene expression analysis using publicly available data of Homeobox family of genes and Tbox genes for which methylation data was also available for in our data set. Although these tumors were relatively hypermethylated at various sites in recurrent tumors versus non-recurrent tumors, they were either upregulated or non-differentially expressed in recurrent tumors. FoldChange p-value FoldChange (1 vs. 0) Column ID (1 vs. 0) (1 vs. 0) (Description) HOXD13 0.000114061 2.06041 1 up vs 0 HOXA7 0.000172107 1.54672 1 up vs 0 HOXB13 0.000356608 1.23524 1 up vs 0 PITX1 0.000571527 3.42517 1 up vs 0 HOXA5 0.000649674 1.56913 1 up vs 0 HOXA1 0.00152652 1.66169 1 up vs 0 HOXD12 0.00329604 1.18036 1 up vs 0 PAX6 0.00709951 1.90785 1 up vs 0 PAX9 0.0174741 1.12557 1 up vs 0 HOXA10 0.0179756 1.23869 1 up vs 0 POU4F1 0.0416546 1.05959 1 up vs 0 HOXC10 0.0477096 1.40787 1 up vs 0 HOXA3 0.0608012 1.18098 1 up vs 0 HOXB3 0.0613335 2.05141 1 up vs 0 FOXA2 0.0674789 1.07005 1 up vs 0 TBX3 0.0701179 1.06385 1 up vs 0 HOXC11 0.0971162 1.10363 1 up vs 0 NKX2-3 0.106482 1.04019 1 up vs 0 HOXB5 0.107014 1.06562 1 up vs 0 HOXB4 0.11069 1.09051 1 up vs 0 IRX3 0.120743 −1.79627 1 down vs 0 HOXC9 0.133088 1.52349 1 up vs 0 HOXD10 0.134932 1.07023 1 up vs 0 HOXD9 0.153315 1.06508 1 up vs 0 PAX2 0.156918 −1.22567 1 down vs 0 HOXB7 0.159887 1.10388 1 up vs 0 HOXA4 0.160191 1.06741 1 up vs 0 HOXB6 0.166413 1.06457 1 up vs 0 NKX2-8 0.245824 −1.05089 1 down vs 0 HOXC6 0.259377 1.4128 1 up vs 0 HOXA13 0.279626 1.06615 1 up vs 0 NKX2-5 0.281508 1.03816 1 up vs 0 OTX2 0.286161 1.02241 1 up vs 0 PBX1 0.293154 −1.09442 1 down vs 0 TBX2 0.31776 −1.11363 1 down vs 0 OTX1 0.350332 1.04444 1 up vs 0 PITX2 0.379015 1.0237 1 up vs 0 POU4F2 0.411171 1.03497 1 up vs 0 NKX6-2 0.411643 1.03463 1 up vs 0 DLX5 0.422143 −1.07733 1 down vs 0 HOXD8 0.433142 −1.06484 1 down vs 0 HOXD1 0.435759 1.03981 1 up vs 0 PAX1 0.463461 1.02538 1 up vs 0 HOXA2 0.468008 1.05235 1 up vs 0 HOXD1 0.479934 −1.01955 1 down vs 0 NKX6-1 0.544333 1.03998 1 up vs 0 HOXD11 0.581698 1.0156 1 up vs 0 HOXA6 0.655808 1.02198 1 up vs 0 HOXB8 0.664046 1.01057 1 up vs 0 HOXC8 0.668768 1.02739 1 up vs 0 DLX6 0.680374 1.01573 1 up vs 0 TBX1 0.693979 1.01974 1 up vs 0 IRX4 0.694484 −1.07894 1 down vs 0 HOXB1 0.703823 1.01884 1 up vs 0 HOXC5 0.767971 −1.03447 1 down vs 0 HOXD3 0.835941 −1.01269 1 down vs 0 PAX3 0.843197 −1.00904 1 down vs 0 NKX2-1 0.865339 1.00688 1 up vs 0 HOXC4 0.868251 1.0286 1 up vs 0 PAX7 0.877734 −1.00675 1 down vs 0 MNX1 0.88141 1.00704 1 up vs 0 HOXC12 0.897218 −1.0118 1 down vs 0 FOXB1 0.913163 1.00606 1 up vs 0 PAX5 0.940595 −1.00921 1 down vs 0 IRX5 0.993078 1.00224 1 up vs 0 Annotation: 1 = recurrent, 0 = non-recurrent

Although preferred embodiments of the invention have been described herein, it will be understood by those skilled in the art that variations may be made thereto without departing from the spirit of the invention or the scope of the appended claims. All documents disclosed herein, including those in the following reference list, are incorporated by reference.

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1. A method of predicting recurrence free survival in a patient with meningioma comprising: (a) determining a tumor DNA methylation profile from a tumor sample from the patient, the tumor DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9; (b) calculating a risk of meningioma recurrence based on comparing the tumor DNA methylation profile and a reference methylation profile comprising the extent to which the methylation status of the at least 200 loci is associated with a risk of recurrence.
 2. The method of claim 1, wherein the at least 200 loci is at least 500 loci, at least 1000 loci or at least 1500 loci.
 3. The method of claim 1, wherein the at least 200 loci is at least 2000 loci, at least 2500 loci, or at least 3500 loci.
 4. The method of claim 1, wherein the at least 200 loci is at least 4000 loci, at least 5000 loci or at least 6000 loci.
 5. The method of claim 1, wherein the at least 200 loci is at least 7000 loci, at least 8000 loci or at least 9000 loci.
 6. The method of claim 1, wherein the at least 200 loci is at least 9500 loci or at least 9529 loci.
 7. The method of claim 1, wherein the calculating the risk is based further on at least one of the WHO grading and Simpson grading of the tumor.
 8. The method of claim 1, further comprising calculating a predicted number of years of recurrence free survival (RFS Score).
 9. The method of claim 8, further comprising stratifying the patient into high, medium and low risk categories based on said RFS Score compared to known reference patients.
 10. The method of claim 9, further comprising treating the patient with adjuvant therapy if the patient is in the high or medium risk category.
 11. The method of claim 10, wherein the adjuvant therapy comprises chemotherapy or radiotherapy.
 12. An array consisting of the 9529 probes set forth in Table
 9. 13. A computer implemented product for predicting recurrence free survival in a patient with meningioma comprising: (a) a means for receiving values corresponding to a tumor DNA methylation profile from a tumor sample from the patient, the DNA methylation profile comprising the methylation status of at least 200 loci represented by the probes set forth in Table 9; (b) a database comprising a reference methylation profile representing the extent to which each said methylation status of the at least 200 loci is associated with a risk of recurrence; and (c) a means for calculating a risk of meningioma recurrence based on a comparison of the tumor DNA methylation profile and the reference methylation profile.
 14. The computer implemented product of claim 13, wherein the computer implemented product further outputs a recommendation for adjuvant therapy if the patient has been classified into a medium or high risk group.
 15. A computer readable medium having stored thereon a data structure for storing the computer implemented product of claim
 13. 